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Full text of "Transactions of the Royal Society of South Australia, Incorporated"

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



Transactions of the 

Royal Society of South 
Australia 

Incorporated 



Contents 



Bayly, I. A. £. Inland-water calanoid copepods of Kangaroo, King and Flinders Islands: 

Biogeography and Ecology -- 1 

Robertson, G. B., Prescott, J. R. & Hutton, J. T. Thermoluminescence dating of volcanic 

activity at Mount Gambier, South Australia ----- 7 

Davies, K. A. & Lloyd, J. Nematodes associated with Diptera in South Australia: a new 
species of Fergusobia Currie from a fergusoninid and a new record 
of Syrphonema Laumond & Lyon from a syrphid - 13 

Campbell, E. M., Twidale, C. R., Hutton, J. T. & Prescott, J. R. Preliminary 

investigations of dunes of the Gawler Ranges province, South Australia 21 

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

Hydryphantidae) --------- 37 

Barker, S. Seventeen new species of Castiarina (Coleoptera: Buprestidae) - 41 

Kolesik, P. A new genus and three new species of Cecidomyiidae (Diptera) from 

Olearia spp. (Asteraceae) in Australia 61 

Brief Communications: 

lyier, M. J., Barrie, D. J. & Walkley, R. W. First fossil record of the hylid frog Litoria 

raniformis (Keferstein) - 69 

Hero, J.-M., Fielding, S. & Retallick, R. The tadpole of Litoria revelata Ingram, Corben 

& Hosmer, 1982 (Anura: Hylidae) ------ 71 

Obituaries: 

Ludbrook, Nelly Hooper MBE, MA, PhD, DIC, FGS. - 74 

Edmonds, Stanley Joe BA, BSc, MSc, PhD, Dip Ed. 78 




PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 



TRANSACTIONS OF THE 

ROYAL SOCIETY 
OF SOUTH AUSTRALIA 



INCORPORATED 



VOL. 120, PART 1 



TRANSACTIONS OF THE 
ROYAL SOCIETY OF SOUTH AUSTRALIA INC 

CONTENTS, VOL. 120, 1996 



PARTS 1 & 2, 31 MAY, 1996 

Bayley, I. A. E. Inland-water calanoid copepods of Kangaroo, King and Flinders Islands: 

Biogeography and Ecology - -_._-. j 

Robertson, G. B M Prescott, J. R. & Hutton, J. T. Thermoluminescence dating of volcanic 

activity at Mount Gambler, South Australia ----- 7 

Davies, K. A. & Lloyd, J. Nematodes associated with Diptera in South Australia: a new 
species of Fergusobia Currie from a fergusoninid and a new record of 
Syrphonema Laumond & Lyon from a syrphid - ■ 13 

Campbell, E. M., Twidale, C. R., Hutton, J. T. & Prescott, J. R. Preliminary investigations 

of dunes of the Gawler Ranges province, South Australia - - 21 

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

Hydryphantidae) ---------- 37 

Barker, S. Seventeen new species of Castiarina (Coleoptera: Buprestidae) 41 

Kolesik, P. A new genus and three new species of Cecidomyiidae (Diptera) from 

Olearia spp. (Asteraceae) in Australia * ■* «■ - 61 

Brief Communications: 

Tyler, M. J., Barrte, D. J. & Walkley, R. W. First fossil record of the hylid frog litoria 

raniformis (Keferstein) - ------- 59 

Hero, J.-M., Fickling, S. & Retallick, R. The tadpole of Litoria revelata Ingram, Corben 

& Hosmer, 1982 (Anura: Hylidae) - - - 71 

Obituaries: 

Ludbrook, Nelly Hooper MBE, MA, PhD, D1C, FGS. - 74 

Edmonds, Stanley Joe BA, BSc, MSc, PhD, Dip Ed. ..... 78 



PARTS 3 & 4, 29 NOVEMBER. 1996 

AnstLs. Mi & Littlejohn, M. .1- The breeding biology of Liioria subgltmtlMiosa and jL film;™ 

(Anura: Hylidae), and a re evaluation of their geographic distribution- 83 

Dyson, I, A. Stratigraphy of me Ncoprotcrozoie Aruhna and Depot Springs subgroups. 

Adelaide gcosyneline - 101 

Dyson, I. A. Stratigraphy of the Neoproterozoic Tent HLI1 Formation and Simmens 

quartzite at South Tent Hill on the- Stuart Shelf, South Australia- - 117 

Taylor, G. S., Austin, A. D. & Davies, K. A. Biology of the eucalypt gall -forming Ry, 
Fergus otiinu Jhiviajrnis MallocK (Diplera: Fergusoninidae) and its 
HHWMNHM'I hymenopterans in South Australia,, wjth a description of a new 
species of Bracon (Hymenoptera; Braconidae ) - 131 

Bird, A* F. Studies On the soil-inhabiting tardigrade, Mmrobiotus cf, pseudohufelandi. 

Iron"! South Australia _ . . . _ _ _ _ 147 

Rolesik, P. Rhopaiomyia goodenkie, a new species of Cccidomyiidac (Diprcra) 

damaging Goodenia tunata (Goodeniaceae) in inland Australia - - 155 

Nicholas, W. L. Robustnema jbsteri sp. BOtt , gen. no 1 *'- (Xyalidac, Manhysterida, Nematoda), 

a common nematode of mangrove mudflats in Australia - - 161 

Smales, L. R. A ^description of Asprrsentis xanvhhrhyrtchi [Johnston & Best, 1937 1 

comb. nov. (Heteracanthocephalidae: Acamhocephala) - - 167 

Brief Communicaiions: 

Baker, G. 11. Seasonal activity of the earthworm, Gemascotex lateralis (McgascolccKlac). 

in a Eucstypoa woodland in South Australia ----- 173 
O'Callaghan* M + G+ & Bcverldge* 1. Gastrointestinal parasites of feral cats in the Northern 

Territory ----------- 175 

Terrace, I. L. & Baker. U. II, Predation of earthworms by the land planarian. Aus(rahpLatui 

sanguined (Moseley) var. alba (Dendy) sensu Jones, 1981 (Tricladida: 
Gcoplanidac) ----------- 177 

Tyler, M. J. & Williams C. R. Mass frog mortality al two localities in Soulh Australia 179 

Irnrn m ZhntMrUnnf cfihr Royal Soaeiy afStmh Auatmlta. \bt i2J. para I A 2, W May. t997 



INLAND- WATER CALANOID COPEPODS 

OF KANGAROO, KING AND FLINDERS ISLANDS: 

BIOGEOGRAPHY AND ECOLOGY 



ByL A. E. Bayly* 



Summary 

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

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



lhin.satthm\ <>J tkf A'ovw/ Sociciy of S. Aust, (1996). 120(1), 1-& 

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

by I A. E. Bayly* 

Summary 

Bayly, I. A, E. (1996) Inland-water calanoid copcpods of Kangaroo, King and Flinders Islands: Biogcography 
.*nd Ecology, ham. R $$£ S. AusL 120(1), l-h, 3! May, 1996- 

Calanoid copcpou* identificaiions arc provided rbr samples from 1ft localities on Kangaroo Island. 18 on King 
Islund und II on Flinders Island. The number of species found was five, llirec and seven, respectively Conductivity 
data are given for most localities, Species richness in relation to land area is tabulated and discussed Bncvkctta 
major is recorded from South Australian territory for the first time The occurrence qf Hcmlbrn-LkvUa .\earli 
iti temporary pools and amongst dense macrophvtes in Iakc c . may be due to the absence ol predators in young 
pools and the difficult? encountered by predators in searching dense weed-beds in lakes, The disjunct distribution 
tit Caiatnnri i<t gibhosa is explicable on the hasis ol easi to west dispersal along a lowland plain during the Pleistocene 
when sea levels were low, followed by marine miudation. 

Kit Woros: Conepoda. Calanoida. biogeography. ecology. 



Introduction 



Methods 



Following ihc glacial (and aridity) maximum that 
occurred in the Late Pleistocene at about 18 ka B.P.. 
deglaciation. and the consequent rise in sea level 
(Chappell 1978; Galloway & Kemp 1481). converted 
several areas of land along the southern margin of the 
Australian comment into islands. The nature and fate 
of ibe samples of the fauna ol greater Australia pnwided 
by these islands is a matter of considerable interest 
Rawlinson (1974) studied th$S issue with reference to 
the reptdes of Bass Strait islands and Tasmania and 
showed that a whole series of Island? in Bass Strait 
and oii' South Australia became isolated during the 
period 16.5 - 63 ka B.P 

Despite their small s»/e and the ability of many of 
Iheni to produce desiccation- resistant eggs, freshwater 
calanoid copepodsare widely acknowledged to exhibtt 
poor dispersal ability (Bayly & Maly 1991; Banarescu 
1990; Bayly 1995). As a consequence, the 
biogeogmphy of calanoid copepods is of considerable 
interest, with dispersal playing a smaller role than has 
Urt n iUppO^ed by the numerous workers who 
simphsueally equate the possession of resting eggs with 
good powers ol dispersal (Bayly & Morton 1978*. 

This paper aims to examine the relationship oi the 
calanoid fauna of three offshore islands (Fig. J) with 
that of mainland Australia and Tasmania and to 
consider Ihc role of historical and ecological laclors 
iu observed differences and similarities. 



* Di:|MrtinentorH(.nli>tiv and Kolultonary Btologv, Mtinash 
I'n'A'T.iiy Clayton Vie 5168; 

Current uddre.vs: 114 Belgruvc Hallain Road litlfwve South 
Vic Slot) 



Racb body of water was thoroughly sampled with 
a zoopiankton net of mesh size 150 /im. Collections 
were preserved in 10% formalin. Conductivity of a 
water sample taken from the field 111 a polyethylene 
bottle was determined in the laboratory with a 
Radiometer CDM2e conductivity meter. Where the 
K-, s exceeded 5.0 mS cm 1 , the conductivity value was 
converted to a salinity value using the method ol 
Williams (1986). For the King Island localities, pH was 
measured with a Meirohm E599 portable pH meter. 

With two exceptions, the Flinders Island localities 
were sampled by the author alone at various limes 
between 1985 198X. and by the author working with 
a lunnologieal team from 10-12 February 1993. The 
King Island samples, with one exception, were taken 
by a team of workers Jnctudine the author during the 
period Z-5 December 1991, With two exceptions, the 
Kangaroo Island samples were collected by the author 
alone during the period 26 June 3 July 1994. 

kt'suhs 

Pliysico-chemical arid biological results are summar- 
ised in Tables 1 - J, Five calanoid species were record 
ed from Kangaroo Island, with only two species, 
Htutkflln rriartimfaia and Cnhnnovna clitetlata. 
occurring at those localities with a salinity of 3.3 g I ' 
or more. At the less saline localities. B. major was 
resmcted to temporary waters. Only three species were 
found on King Island, and one ol these, B. pscudo- 
chelae, occurred in the sole temporary warer body that 
was sampled. C taMtianieu was the only species Ibund 
in waters with a recorded pH of less than 6.0. None 
of the King Island waters included in the survey was 
saline. Seven species were recorded from Flinders 
Island with C hwmrtiut most common, B. symmetrica 
was the only species common to all three islands. 



2 I. A. E. BAYLY 

Table 1. Occurrence of calanoid copepods on Kangaroo Island. 



Locality 



Sampling 
date 



Permanency* 



Conductivity 

(mS cm' 1 ) 



Salinity 
(g I 1 ) 



Bm 



Species* 1 
Bs Bt 



Ca Cc 



Dam 1 near Penneshaw c 
Dam 2 near Penneshaw c 
Waterhole edge Edwards Lagoon 
Pond Roper Road 
Lake at Karatta 
Pond nr rush Lagoon 
Pond south end Roper Road 
Ditch east Kingscote Airport 
Small Grassdale Lagoon 
Big Grassdale Lagoon 
Kaiwarra Cottage Pond 
Duck Lagoon 
Discovery Lagoon 
Lake Ada 
Murray Lagoon 
White Lagoon 



20.viii.1991 


P 








X 




20.viii.1991 


P 








X 




30.vi.1994 


T 


0.21 




X 






l.vii.1994 


T 


0.22 
0.31 




X 


X 




28.vi.1994 


SP 






X 




27.vi.1994 


T 


0.44 




X 






l.vii.1994 


SP 


1.23 






X 




27.vi.1994 


T 


1 71 




X 


X 




28.vi.1994 


SP 


3.00 






X 




28.vi.1994 


P 


3.65 






X 




2.vii.l994 


T 


5.8 


3.2 


X 


X 




l.vii.1994 


P 


6.0 


3.3 






X 


27.vi.1994 


T 


12.8 


5.J 






X 


3.vii.l994 


P 


7.5 






X 


3.vii.l994 


P 


15.3 


9.1 






x 


27.vi.1994 


SP 


153 


51.3 









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

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

c. collected by N. Frick 




Kangaroo Is.C^r^ 





King Is. 



^} Flinders Is, 



<^ 




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



CALANOIDS OF KANGAROO, KING AND FLINDERS ISLANDS 



Tabi p. 2. Occurrence of calanoid copepods on King Island. 



Locality 


Sampling 


Permanency' 


Conductivity 


pH 


Species* 




date 




(mS cm"') 




Bp Bs Ct 


Pool m Currier 


8.xi l%3 


T 


- 




X 


Meatsafe Lagoon 


5.xb.l99l 


P 


0.56 


3.8 


X 


Dead Sea 


2.xii 1991 


P 


0.56 


4.7 


K 


Lake Martha Lavinia 


3.xit.l991 


P 


0.75 


5.2 


X 


Seal Rock. Lagoon (North) 


4.xii 1991 


P 


0.96 
1.04 


7.9 


X 


Pearshape Lagoon 


2-.\ii 1991 


P 


7.8 


X 


Lake nr Surprise Bay Homestead 


4.xii.l99l 


SP 


UO 


7.4 


X 


Lake opp. Pearshape Lagoon 


2.xiU99l 


P 




8.2 


X 


Pennvs Lagoon 


3.xii.l99l 


P 


1.15 


7.7 


X X 


Granite Lagoon 


5-Xii 1991 


P 


1.26 


6.4 


X 


Pioneer Lagoon 


4.xii.l99l 


P 


1.46 
1.47 


7.8 


X 


Lake btn Denbys & Pioneer Lagoon 


2.xii.l991 


P 


7.2 


X 


Denbys Lagoon 


2.xii 1991 


P 


1.57 


6.6 


X X 


Lake cast end Pioneer Utgpdn 


4.X1.J991 


sv 


S 


8.5 


X 


Seal Rock Lagoon (Middle) 


4.xii.l99l 


SP 


7.5 


X X 


Cask I^ke 


3.xii.l991 


p 


L96 


8.2 


X 


Lake Wickham 


3.xii.l991 


SP 


2.15 


8.9 


X 


Lake Flannigan 


3.xii.l99l 


SP 


2.28 


9.9 


X 



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

b. Bp - Boecketla pseudochelae Searle; Bs - K symmetrica Sars; Ct = Catarwrcta tasmanica (Smith); 

c. collected by M. J. Littlejohn. 



Table 3. Occurrence of calanoid copepttds on Flinders Island, 



Locality 


Sampling 


Permanency 


Conductivity 


Salinity 


Species 


b 








date 




ig, 


is n 


Bm Bp Bs 


tit 


Hs C 








(mS cm' 1 ) 












Brodies Lagoon 4 " 


May 1962 


T 








X 






Scotts Lagoon c 


May 1962 


T 












X 


Pond nr Sticks Lagoon 


15.V.1985 


T 












1 


Pond (t) nr Kilheerankie 


5.VI.1988 


T 






X 








Pond (2) nr Killiecrunkie 


S.vi.1988 


T 






X 








Reedy Lagoon 


9.vi 1988 


P 






X 






* 


Shag Lagoon 


12. ii. 1993 


T 


1.7 










X 


Lake btn N & S Patriarchs 


J9.V.1985 


SP 


2.2 








X 


X 


Small lake (1) 


















nr Singleton's Lagoon 


I0.ii.1993 


T 


2.4 










X 


Small lake (2) 


















nr Singleton's Lagoon 


10. ii, 1993 


T 


5.1 


2.7 








X 


Sticks Lagoon 


15.vvl985 


T 


12.3 


71 




X 







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

b. Bm - Boeckella major Scarle; Bp = B. propinqm Sars; Bs = B. symmetrica Sars; Bt = B. triarticulata (Thomson); 
Hs = Hemiboeckelta searli Sars; Cg = Calamoe.ia pjhbosa (Brehm); Ct — C tasmanica (Smith). 

c. collected by W. D. Williams 



Discussion 

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



are the only exceptions) despite wide fluctuations in 
population density (Bayly 8c Williams 1973). In any 
large sample at least some males and egg - or sperma- 
tophore-beanng females are present and the species 
determinable. This means that yearly or seasonal 
differences in sampling dates should not unduly 
intluence the assessment of 'Jte fauna of the per manent 
waters. This leaves a residuum of problems for 
comparisons which, however, arc not so great as to 
preclude the examination of a number of general 
features and trends. 
The much cited island biogeography theory ot 



I. A t, BAMy 



MacArtiiur A Wilson \\%7) would predict thai a 
[>om'iyc correlation should exist beiween the number 
ol if^tctes found within a discrete urea and the size of 
thiu area. With respect to non-marine calanoids in the 
Australasian region. Table 4 indicate? thai across the 
whole spectrum of su land masses there is only a very 
rough correlation of the sort predicted. Several 
anomalies call fOt consideration and explanation 

King Island and Flinders Island differ only slightly 
in area bui the former apparently ha* less than hah 
die number of speue.-- found on the latter This 
diffcicnee, if it iv not an artefact is difficult tn explain 
but it may be significant that native habitat destruction, 
including the removal of vegetation, has pmceeded i< 
a greater extent an King Island than Flinders Island 

Kangaroo Island is about three and a half limes larger 
than Flinders islaruJ but has fewer calanoids (if the 
haiobiont species C chwllata is omitted it lias only 
four species), it is difficult nowadays to bud a large 
natural body of fresh water on Kangaroo Kland, 
Extensive vegetation removal in the period 1045 - 1955 
andtheeonsequenl rise in water tables and mobilisation 
of salt has resulted in the saJiriisation of several lowland 
lakes thai were formerly fresh. Murray Lagoon 
originally contained fresh water but today it is saline 
(salinity 91 g I ' on 3 July 1994; Table I). Several of 
ilv litkes on Kangaroo Island visited during the winter 
i'A I994 were found to be highly saline and were noi 
sampled lor that reason. Half of the natural fresh waters 
that were located were very small and iempora»y '" 
charadei Jt is conceivable that species like Caiamoecia 
gihboMi and C, lasmunuo, which occur in south- 
eastern and south -western Australia and typically 
inhabit permanent fresh waters, no longer oecui on 
Kangaroo Island as a result of recent salinisaiion 

New Zealand is about four times larger than 
Tasmania but ha* fewer ealaiioids. However, during 
the Ohgocene. vmic two-thirds of the area of modern 
New Zealand was covered by sea (Stevens 19X0) 

Roeckelta symmetrica., which occurred on all tho-c 
IslOfKfc. and B. trianictdata, which wa-> found on 
Kangaroo and Hinders Islands, are both common and 



Iahii 4, S^ml nun and specie* nehnexx 



Name id land mass 



4ttA 

tki"-) 


Nimibi 
ruliifiuiJ s 


1.200 
1,330 

4,400 

67800 
2nv>.000 

7.682,000 


3 

7 

H 

m: 
*3 



King (slam) 
Flinders Islam! 
kdftgaiuo IsMtuJ 
TaMrtjnui 
New Zealand 
AuMralta 



a. Belonging io ilie tatmly Ccntronagiilae and tunneled lit 
the genera B&eekello. Hemiboftkrtla and G/lamoecia. 



widely distributed species (Bayly J992a). The 
occurrence of B. propinqua on Flinders Island only 
(Table 3i is consistent with the existing evidence lhai 
within Australia, this species is restricted to the fa* 
eastern fringe of the continent, previous Australian 
records are from the east coast of Tasmania and coastal 
New South Wales 8. major (Kangaroo and Hiudcis 
Islands; Tables I and J) is characteristic of temporal;, 
waters and has been recorded previously from New 
South Wales. Victoria and Tasmania (Bayly 1992a! 
Tire present record from Kangaroo Island is the first 
from South Australian territory hut \\ is likely that itlii 
merely reflects a lack of investigation in Ibis Stale of 
ihe copepocb of temporary ponds and pools. ft 
pscudochelof (King Island: Table 2) is another 
Icmporaiy water specialist previously noted from 
southern New South Wales, Victoria and Tasmania 
Odamovaa uuimmiai (King and flinders Islands) and 
C Gtflplftla (kangaroo [stand) are widely distributed 
species known from south-eastern and south -western 
Australia (Bayly 1992a). 

HctniOocxkt'Ita wurlt typically occurs in temporary 
ponds and pools, but, as with the present record u..n. 
Flinders Island (Table 3), it also occurs in littoral weed 
beds in permanent or semi-permanent watery This 
commonality of occurrence is not as incongruous as 
il first appears. Water permeating dense vegetation fa 
the littoral region of a permanent lake has an ecological 
similarity |0 thai in a shallow, temporary pool 
(including those entirely devoid of vegetation! thai i$ 
not commonly appreciated namely the exclusion ol 
piv-datofv Jt in well appreciated thai, in a newly formed 
pool, flying insect predators such as notoneetids may 
take some time to arrive and, until this occurs, ilu* 
habitat may be largely predator-free. The fact that 
mtnibot'ckella characteristically occurs very early in 
pool successions (Bayly 1992b) suggests a high degree 
Of predator susceptibility. However, a.s pointed out by 
Connell (1975) some prey species have evolved the 
ahdity to live in refuges that the predator cannot invade 
because the habitat smuturt' is raodiffiru/r f0 Atv/rr/i 
It is presumably for this reason that dense littoral 
vegetation oilers H searh a refuse from limnetic fish 
and insect predators in lakes. H uarii iv widel> 
distributed, occurring in south-eastern and south- 
western Australia 

Cahtmuffa xtbbosa has the most intriguing 
distribution of all the Australian freshwater calanotds, 
it occurs in Tasmania, on Flinders Island, along the 
coastal fnngc of south-eastern Sotftfl Australia between 
Ml Gambier and Salt Creek and on two gram ie 
outcrops near Balladoma in Western Australia (Bayly 
1984. 1992a) A previously unpublished record is from 
near Mr Rough to the south cast of Salt Creek in South 
Australia. The two Western Australian populations 
were treated (Bayly 1979, 1992a) as belonging to a 
separate subspecies from the eastern form. 



CALANOIDS OV KANGAROO, KING AND FLINDERS ISLANDS 



The most reasonable explanation for current 
disjunctions m the distribution Of C i>ihht>.\d is thai, 
at the lime of one of the three glacial maxima during 
the L ate Pleistocene iChappell 1978). probably the last 
one at 18 ka B P, it was conlirmously distributed ;tlong 
a coastal lowland plain to the south of the present 
southern coastline of Australia. It may be supposed 
to have extended from eastern Bass Strait to the western 
limit of the Great Australian Bight kf Nelson 1981, 
rlj: 2 ) -A subsequent rise in sea level of more than 
100 in iChappell l°7S) would then have been 
responsible tor the present day disjunctions. The 
morphological ev'tdeuce suggests that the Western 
Australian form is a derived rather ihan ancestral form. 
Thus it h proposed that, some time widtin the 
Wisconsin glaciation when *ea levels had been lowered 
hy about 100 m. gradual east to west dispersal of C 
glbbttMt occurred along a broad coastal plain that is 
now submerged. This proposal of east lo west 
dispersal, followed by subspeeiation in the west as n 
result o1 vicanant isolation, parallels the pattern of 
specialion in Western Australian frogs first proposed 
by Main et ol. (1958) and subsequently adopted by 
Mam (1968) and Little John (1981). It should be noted, 
however, that more recent molecular data on frogs is 
said acrt |o support multiple east lo west invasions 
during the Pleistocene as being the explanation lot 
specialion in Western Australia (Roberts & Maxson 

I'-JXY, 



If we accept submergence of the southern plain as 
the explanation for the disjunctions in ihe distribution 
of C gibbosity then three explanations may be offered 
for the apparent absence o! this species from Kangaroo 
Island and King Island. First, the species does occui 
on these islands bui the present samplings were noi 
intensive enough to reveal it. Second, C. gtbbosu war 
originally present on ibese islands bui subsequent 
ecological changes (e.g. salinisaiion on Kangaroo 
Island) have brought about its local extinction. Third, 
although the original distribution 0fCgfb$O$Q Bfoflg 
the now submerged plain was continuous in a broad 
sense, it was nevertheless somewhat patchy, and the 
persistent land sample* provided by these two islands 
were nol sufficiently Jaige to include ibis cuibnoul. 



AckitowliUfcincnts 

During the field worit on King island in December 
\ l )9\, I was accompanied by Russell Shiel. Grace and 
Peter Tyler and Robert Walsh. On Flinders Island in 
February 1993 1 had the company of Peter Kew. Colin 
Magillon, Russell Shiel , Peter Tyler and Robert Walsh 
1 thank alt of these people for their companionship and 
assistance. The field work carried out on King Island 
and Flinders Island was supported by a National Estate 
Grant to P. A. Tyler. Additionally. Monash University 
COt1lTibmed funds towards the King Island expedition 



References 



BANAktfscu. P (1990) -Zoogeography ol fresh waters, Vol. 
1. Genera) diMrihuiinn and dispersal ol freshwater animals" 
(Al'LAVctlag. Wic^baden). 

Bavi v. I A. F. irWi Funlier contributions tu a knowledge 
of thecenlropagid genera Boerketh. Hemibtwfo'lhi ami 
( V«/r/w*»t(wtaihiljisu-calamudeopept.x1s). Aust t. Mat. 
Frvtfw Res M\, |0>127 

(1984) A. new species trf CaLtmoeain (Copepwla: 

Calanoida) from South Australia, and comments on three 
congeners. Trans. R. Sfcc 5 -4m v/. IUX, 147 154. 

_ 0992a) -The Non-marine Cenimpugidae (Copept^a: 
Calanoidal of Ihc WbrKT (Gulden lo flic Identification of 
the micnnnvertcbrates- of the com mental water, ol Uic World 
No, 2| fSPB Academic Pubbshmg, The Hague). 

(1992b) The roic-ro-Crusucea iwl physicu chemical 

feature,* ol temporary ponds near Nortricliite Western 
Australia, l /?. S<k. W Ami. 75, <*M06. 

( 1995) Distinctive aspects of thf ;i <oplankton of large 

lake* in Australasia, Anurclka and Soulb America Mur. 
Freshwater Res. 45, JIO9-I120. 

tV Mai.v, E. i 1 1991) Copepod conundrums, reply 
lo Jefferson T Turner / Bio^aer 1M 468-469. 

& Mobtok. ft W. (19781 Aspects ol rhe 



/<M)ge<i«rapbv of Australian microcrusiaccftii*; Verb, (nt 

Veretn. 7Wr. Arisen. Uttmtti 20, 2537 -3540. 
& Williams, W. D. <I*)73> "inland Waiers and their 

lieology" (Longman, Melbourne). 
Chapheli., J- (1978) Chronological methtxls and ilie ranges 

and rales of Quaternary physical changes pp. 1-34 fn Walker, 



O. & Guppy, J. C. (Hdr.i ^Biologs and Quaternary 
Environment" tAusr Axad .Sci., Canberra!. 

Connh-I-, J. H. (1975) Some mechanism* producing 
Structure m natural eommunriies; a model ami evident*** 
from field experiments pp. 46tt-490 hi Only, M, I. i: 
Diamond, J. M. (Edi) 'rcoloj*y and b'vollilion of 
Communities' --Harvard Umv Pies:.. Cambridge"). 

G/vu<dwa>\ R. W. A KtMi-, ti, M. H98J) Late Cain<»/iMc 
envirwimeiits in Australia pp. 52 -St) hi Keasi, A. (Od ) 
■Ecological BicigCi'graphy rf Austral*** Vol I (Junk, The 
Hague). 

Littlejohn. M. .1, (19^11 The Amphibia <>V »u-ku M»mherrj 
Aiisiralia i /»x>t;eot;uphic r^rsj»cctive pp I3D5-B3U, tbui. 
Val || 

MVCARTII^i, K. II. & WlJLSON. t. O *N67) "The Theory 

<A Island Bioet.v>:raf»hy" jMcnographs in population Unlugy 

Ni>. 11 (fTincetcni Univ. Press, Princeton). 
Main. A. R. (1968) Eculogy. systenurics and evolution ot 

Australian frogs itivtmc hcol. fljpy. 5, 37 86. 
, LEE, A. K & LirrLCJOHN. M J (1958) Evohilion 

in three genera of Australian frogs, SfoJlMffcM U. 7l± 2} 5 
Nri-SO\, E. C. (1981) Phyiogtxijrraphy of'soulhern AuMraJu 

pp. 7*5-759 in K-.ast. A. {F.d.) "Ho>logical Bn>geot;ruphv 

of Ausunlia" Vol I (Junk* The Hague). 
Raw i iNSf>N, P A. i\*¥74\ Bio^eography and ecology ol ibe 

reptiles of Tasmania and the Bas> Strait area pp. 29I-33H 

h Williaitis, W. P (Kd.) "Bio^eography and ecology ol 
lasnuinia' (Junk. The Hague), 



6 I. A. E. BAYLY 

Roberts, J. D. & Maxson, L. R. (1985) The biogeography Stevens, G. R. (1980) "New Zealand Adrift" (A. H. & A. 

of southern Australian frogs: molecular data reject multiple W. Reed, Wellington), 
invasion and Pleistocene divergence models pp. 83-89 In 

Grigg, G., Shine, R. & Ehmann, H. (Eds) "Biology of Williams, W. D. (1986) Conductivity and salinity of 

Australian Frogs and Reptiles" (Surrey Beatty, Sydney). Australian salt lakes. Aust. J. Mar. Freshw. Res. 37, 177-182. 



THERMOLUMINESCENCE DATING OF VOLCANIC ACTIVITY 
AT MOUNT GAMBIER, SOUTH AUSTRALIA 



ByG. B. Robertson*, J. R. Prescott* & J. T. HurroNf 



Summary 

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

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



TrcmMVi'ums oftht- fowl Sovitty of'S. Ausi (1<W6). 120(1). 7-12. 

thermoluminrs(t.nc:k dating of voijcaxic activity 
at mount gambier, solth australia 

by G. B. Robertson* J. R. Presoott* & J. T. HurroM 

Summary 

RoBURreoN, G. B, T Presto-it, J. R. 8l Hutton. J. T. (1996) Thermoluminescence dating of volcanic activity 
;ii Mount Gambier, South Australia. Tains. R. .W«. S. Aust, 120(1), 7-12, 31 May. IWfi. 

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

KtY Words. Thermoluminesccnce dating, volcarrism. Holoeene, Mount Gambier. 



Introduction 

The a* hOJc he*en a number of" attempt? at determining 
the sequence of events and the time scale involved in 
Ihe formation of the volcanic complex at Mount 
Gambier in South Australia's South-east The finding's 
indicate that the volcano as wc sec it today is the result 
of a very complicated series of events, possibly spread 
over several thousand years- The earliest radiocarbon 
dates obtained by Fergusson and Rafter (19>7) indicated 
two main periods of eruption, one at 4700*70 years 
B. P . the other at 1410 ± 90 years B. P. These dates were 
incorporated into a geological history of the eruptions 
by Sheard (1078). Subsequently Blackburn eral (1982) 
concluded that the most likely C-14 age was about 4 
thousand years tka) although charcoal samples were 
found covering the range 3.5 to 8 ka. Barton and 
MclZlhinney (1980) suggested that Mount Gambier must 
pre-date 5-6 ka B.P Recently published work by Leaney 
ft al (1995) on C-14 in the inorganic and organic 
Carboil fractions of Blue Lake sediment cores point lei 
volcanic activity at about 7 ka On the other band 
Shear tl (1995) now interprets the various C-H ages as 
indicating a period u( activity commencing 5-4 3 ka 
B.P. and extending over perhaps 300 years. Nearby 
Mount Scbank has been dated by thermoluminescentc 
at 4.9 i-0 5 ka (Smith & Prescott 1987) in agiccraciu 
wiUt a palacomagneuc measurement of Barbell t and 
Sheanl (19KI) who placed the eruptions of Mount Schank 
and Mount Gambier either between 1 and 5 kn or older 
than 7 ka. 

in 1987, in collaboration with CSTRO. we cmbai ked 



* Doptfituu-iil or Physics and Mathematical Physic^ 

University of Adelaide S, Au*t. 5*105 
t Deceased 



on a programme to date the eruptions at Mount 
Gambier using thermoluminesccnce. This technique 
depends on the measurement of the energy imparted 
to a mineral crystal over time by the ionizing radiation 
generated by radioactive elements in the environment 
and by cosmic rays; this energy is released as lighi 
when the mineral is heated in the laboratory; Its success 
as a dating method depends on the fact that the TL 
was set to zero by the event being dated, in this ease 
either by the heat generated by the volcanic eruption 
or by the bleaching by sunlight of ejected material 
In the lirst instance samples were obtained from sites 
where it was considered likely that sufficient heating 
had taken place to reset the TL clock* and where there 
were likely to be sufficient quantities of quartz for an 
analysis. The quart? is derived from country rock 
sediments through which the volcanic conduit passed. 
The sites chosen were at Valley Lake, at Brownes Lake 
and at the Blue Lake crater, where there were layers 
of heated material underlying the la\a flow. Later the 
scope was extended to include material which may have 
been blown into the air during the event, heated and/or 
bleached during transport and deposited in positions 
more or less remote from the central crater. Tuff 
samples were collected from several sites in the Mount 
Gambier and Mount Schank areas where it was thought 
that (hey might have been associated wilh the lotinatton 
of either of the volcanoes. 



Details of Samples 

Details of the samples collected from the Mount 
Gambier complex are given m Table 1. The mineral 
quoied refers to the material thai was extracted for the 
TT measurements The quart? samples were collected 
and prepared according lo standard practice (Huntley 



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



Table I . Details of the samples collected in the Mount Gambler area. 











TL 


Dale 


Sample 


Site 
Bmwnc& Lake 


Lilhology sampled 




Mineral 


Y/N 


MGIb/3 


Spatter lava 




Quartz 


N 


MG21/I 


Blue Lake pump house 


Healed tuff deposits 




Quartz 


Y 


MG2S/I 


Blue Lake pump house 


Bridgewater Fm. sand below tuff and lava 


Quart? 


Y 


MG2b/|fl 


Blue Lake pulup house 


Hani tuff 10 cm below lava 




Quart/ 
Fine grams 


Y 

N 


MG2c/l2 


Blue Lake cliff behind pump house 


Upper luff (c. 20 m above level of MO 


Quarks 


V 










Fine grains 


N 


M024/J2 


Blue Lake carpark 


Banded upper tuff (same level xs 


MGJc/LI) 


Quart/ 
Fine grains 


Y 

N 


MG-I 


Devil's Punchbowl 


Scdimcnl/lulT 




Quart/ 


N 


MG5S/0.I 


Valley Lake, Nurses Landing 


Baked tuff below basalt (0.1 m) 




Quart/ 


V 


MG5S/0.5 


Valley Lake Nurses Landing 


Baked lull below banaK (0.3 Mi) 




Quart/ 

OJats 


Y 

N 


MG5S/I.5 


Valley Lake, Nurses Landing 


Baked luff below basalt (1.5 m) 




Quart/. 


Y 


MG5 lava 


Valley Lake. Nurses Landing 


Basalt 




Quartz 


N 


Mbesww 


2 km HtMJlh "t Mounl Gam bier 


Tutf, sunlight hleaehed 




Quart/ 


V 


ML, 7 


Pollers Point Lookout 


Ropey lava from path below lank 




Quart/ 


N 


SCIOS/0 6 


Mount Schank 


Hani tuft layer 




Quart/ 


Y 


SCJ2S/a 


MOimt Schank 


H;ml layer <>| bedded luff 




Quart? 


Y 



el ul. 1993), Leading to extraction of 90-125 fim quartz 
grains, the yield amounting to I to 2% of the bulk dry 
weight. Fine grains containing a mixture of minerals 
were extracted from die 4-1 i pni fraction. 

The location of the sites is shown in Fig. 1. 

The spader lava at MG11V3 is described by Sheard 
(1978) as representing the last evidence of volcanic 
activity. Unfortunately no material suitable for TL 
dating was extracted from it. The same outcome 
resulted from attempts to extract quart/ from the ropey 
lava at MG7. 

The MG2 sue at the Blue Lake was extensively 
sampled from the sides of the crater just below the 
pump house where there are heated tuff layers covering 
sands of the Bridgewater Formation and from the cliff 
face above and behind die pump house where the tuff 
layers were deposited as a resuk of fall out from the 
eruptions. 

The group of samples (MG4) collected near the 
Devil's Punchbowl contained terrestrial sediments of 
the Wangerrip Group from below the Gambler 
Limestone, dispersed in the volcanic tuff where it had 
been carried by the eruption. They were found not to 
have been sufficiently heated to make TL 
measurements. 

The MG5S samples from Valley Lake were Collected 
from the baked tuff at 0.1. 0.3 and 1.5 m below the 
base of the lava flow. It was nol expected that the lowest 
level would have been sufficiently heated but it was 
included to check this point. A small quantity of 
volcanic glass was extracted from the MG5S/0.3 
sample but this was highly magnetic and produced an 
insignificant TL signal. Basalt was also collected from 
this site. It contained very small quantities of extracted 




■ 



<0 I1AVM ■ 



Vi ^ ^ 



if# i . 



Wrttm 




I 



-,r p i i r ■ - ,,,,. .... I,, , 



i I ' 



lU't 




Fig. L Lower South east of South Australia showing the 
locations of the volcanic deposits and details of the sampling 
sitej> Adapted from Sheard (1978. 1983). 



TL DATING AT MOUNT GAMBIER 



quuru, and SOO mm sections eul ftoin the bulk sample, 
which might contain other minerals, yielded only low 
levels of TL. 

Other samples of tuff and sand were collected from 
a sue some distance to the south of the lake (MGoS/60) 
and from sites in the Mount Sehunk area (SC serie* 1 , 
I km east of Mount Scharik- 



TL Analysis 

The measurements required in order lo calculate an 
age aie the intensity of the TL tn the natural sample 
and the TL sensitivity, as shown by the TL generated 
by a known amount of radiation. These two values are 
combined lo determine the amount oi radiation dose 
received by Ihe natural sample, the equivalent dose 
It is also necessary to measure the environmental dose 
rate received by the sample in the local environment 
The age is then given by 

Equivalent dose (Gy) 
4B**M- Dose rale (Gyka 1 ) 

where (he unit of radiation dose is the gray (Gy). 

The TL analysis protocol was selected according l»« 
whether the- samples were thought to have been zeroed 
by heat tsamptcs in the MG2 and MG5 series) or by 
sunlight (samples from MG6 and the SC series). In 
the case of the heated samples the usual procedure for 
pottery dating was followed ( AUken 1985). In the case 
of the sunlight bleached samples the selective bleach 
method was used (Prescott & Mojarrabi 1993 V. the 
analysis to give the equivalent dose followed the 
"Australian slide" method (Prescotl et at 1993) 



Dose Kale 

The sample dose rates "were determined by thick 
source alpha counting (TSAC), by neutron activation 
analysis (NAAj. by X-ray fluorescence spectrometry 
(XRS) and/or bv gammy-ray seintillometry ueinh. as 
appropriate lor the estimation kA' the element* urumum, 
thorium and potassium (Prescolt & Hution |SM) Tl>e 
concentrations of these elements arc given m Table 2 
together with the calculated dose rates. The samples 
are grouped according to type and location and it is 
noticeable that the baked tuft below the layer of lava 
has higher concentrations of all the elements than the 
tuff found in other locations and also the sand from 
the Bridgewater Formation Consequently the dose 
rates in the baked tut] are about twice those in >Hn 
tuff samples and higher levels of TL might be expected 
m these. 

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



analysis used, implying that the members of the U and 
Th decay chains are in equilibrium in these samples. 
The worst case is MG2S/1 which shows evidence <»f 
higher values for :Ah U obtained by TSAC and 
scmtillometry compared with Ihe values obtained by 
DNA and by XRS. This situation has been shown lo 
arise in cases of disequilibrium between 'he parent 

**U and its immediate decay products down to ' U U 
and the remainder of the decay chain from - ?0 Th lo 
-'"'Po occurring in samples collected in D similar 
environment at Mount Schank (Smith & Prcsentl 

1987'L The dose rates have been calculated to allow 
lor the deficit in the early part of the chain, taking the 
extreme possibility of a value of 0.80 /<g g 1 for 
aqjdttU and i.4*5 ^g g ' for ' n "Th onwards. 

The dose rates were derived from the element 
concentrations using the conversion factors of Nambi 
and Aitkeu (1986). The values quoted in Table 2 include 
a cosmic ray contribution of 0.10 io 0.12 Gy ka ' 
depending on the site (Prescott & Hiitton 1994). The 
weighted means of the various estimates of dose rate- 
were used in the calculation ol the ages of the samples- 
Age Determinations 

As is* usual for hcat-?craed samples, fust- and 
second-glow growth curves were obtained and the two 
intercepts on the dose axis combined to obtain the 
equivalent dose (Ailken 1985). The growth curves were 
fitted to a linear relationship and used to generate the 
equivalent dose plateau which is shown superimposed 
on the natural glow curve of MG2S/1 in Fig. 2. Valid 




Terrrjerquire {%) 

Fig. 2. Equivalent dose plateau for the MG2S/1 sample with 
its TL glow curve *upermi posed . 




.'■■■ r 



J0U 3*5 3§q J75 

Temperature r°£ ) 



u 00 



Fig. 3. Equivalent dose plateau for the MG5S/03 sample with 
its TL glow curve superimposed. The double plaleaU 
indicates two periods of healing. 



in 



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



TABLfc 2. Concent rations of uranium, thorium ami potassium in the samples, determined by thick-source alpha cou/itiny 
(TSAC), X-ray fluorescence spectrometry (XRS), neutron activation analysts (NAA), delayed neutron analysts (DNA) and 
in situ %um/rui-ray \cintillometry (Scint), together with the resulting dose rates. 



Sample 


Melhitd 


Uranium 


Thorium 


Po labium 


Dose Rate 






M g' 


/^g* 1 


% 


Gy ka < 


Baked tuff below lava 










MG2t/l 


TSAC 


2.34 ±0.48 


6.43 ±1.61 




2.45 ±014 




XRS 


2.60 ±0.40 


8.50 ±0.50 


125±0.0l 


2.67 ±0.10 


MG5S/0.I 


TSAC 


2.71 ±0.27 


8.04 ±0.92 




3.08 ±0.1 




XRS 


3.00 ±0.40 


10.20±0.50 


1.82 ±0.01 


3.32 ±0.1 


MG5S/0.3 


TSAC 


2.46 ±0.41 


10.79 ±1.38 




3,52 ±0,12 




DNA, NAA 


220±0.ll 


9.00 ±0.45 




3.38 ±0.06 




XRS 


2.30 ±0.40 


10.40 ±0 50 


2. 19 ±0.01 


3 51 ±0.08 




Sc.nt. 


2 62 ±0,35 


8.52 ±0.05 


2.03 ±0.05 


3.47 ±0.09 


MG5S/1.5 


TSAC 
DNA 
XRS 


2.0±0.3 
1.87 ±0.08 


69±I4 


1.59 ±0.01 


2.61 ±0.1 


MG5/Lava 


TSAC 


1.93 ±035 


4.59 ± I 15 




2.52 ±0.08 




DNA, NAA 


1.10 ±0.40 


5.70 ±0.50 




2.40 ±0.08 




XRS 






t.50±0.01 




MG2b/K> 


TSAC 
DNA 


2.58 ±038 
2.43 ±0.09 


8.49 ±1.05 








XRS 






2.09 ±0.02 


3.26 ±0.22 


Bridgewatcr 


Formation sand 










MG2S/I 


TSAC (a) 


l.44±0.2l 


4.59 ±0.69 




1,58 ±0.07 




<b) 


1.07 ±0.20 


6.66 ±0.65 




1.64 ± 0.1/7 




DNA. NAA 


0.80*0.10 


5.50 ±0.40 




1.56 ±0.05 




XRS 


0.90 ±040 


7.40 ±0.50 


0.60 ±0.01 


1.48 ±0.08 




Scint 


1.49 ± 0. IK 


5.77 ±0.24 


0.78 ±002 


1.73 ±0.05 


Tuff various 


locations 










MG2e/l2 


TSAC 


J. 41 ±0.26 


3.03 ±0.68 








XRS 






0.66 ±0.01 


1.27 ±0.08 




Scinl. 


1 50±0.12 


4 32 ±0.20 


0.71 ±0.02 


1,51 ±0.05 


MG2J/12 


TSAC 


1.05 ±0.40 


632 ±1.36 








XRS 






0.74±002 


l.5O±0.14 


MG6S/60 


TSAC 


l.90±0.46 


4.70±1.70 




1.66 ±0.13 




DNA, NAA 


1.84 ±0.19 


6.24*0.19 




1.76 ±0.07 




XRS 






0.74±0.05 




SC10S/0A 


TSAC 


2.04 ±0.33 


5.11 ±J. 15 




1.93 ±0.14 




DNA, NAA 


2.32 ±0.60 


7 33 ±0.17 




2.15 ±0.15 




XRS 


1.70±at7 


7.70 ±0.77 


0.94 ±0.10 


2.03 ±0.12 


SCl2S/a 


TSAC 


1.80 ±0.39 


4.06 ±1.33 




I 71 ±0.12 




DNA, NAA 


0.89 ±0.42 


5.26±0.15 




1.58 ±0.10 




XRS 


1.60±0.I6 


6,70 ±0.67 


0.87 ±0.10 


186±0.09 



dating requires that there should be a distinct plateau 
over a range of temperature from about 300° C to 
400°C fWintle & Huntley 1982). This condition is well 
met in the case of sample MG2S/I. Sample MG5S/0.3 
on the other hand, has two plateaux (Fig 3). The 
double plateau suggests that this sample experienced 
two heating events, the second one of which did not 
reach a sufficiently high temperature tD zero the TL 
above 350 ,:I C. Il therefore yields two ages, for differing 
events. Analysis of the phenomenon of the double 
plateau is given in more detail in Robertson et al. 
(1996). The calculated ages are given in Table 3 
together with the equivalent doses and the dose rales 
The analysis of MG2b/10 was not completed because 
there was not enough pure coarse-grained quartz. 



Discussion 

The ages obtained show a spread of values which 
are not easy to interpret but it is interesting to note that 
in the MG samples there is evidence of events having 
occurred at about 4 ka and at about 7-8 ka, coincident 
with other methods of dating. The figures for MG5S 
show the single age for the 0.1 m sample and the two 
ages resulting from the double plateau for the 0.3 m 
sample. 11 was expected that, as the last heating raised 
the temperature of the 0.3 m level to no more than 
350°C, the age of the 1.5 m level would be at least 
7 ka, but the very much older age (98 ± 15 ka) does 
not fit in with the observatioas and the geological inter- 
pretations (Sheard 1978, 1995). It suggests that at least 



ri, IXOTNG AT MOUNT GAMBIFR 



II 



Table J, Ape rstimaws oftk* sampic\ obtained from the equivalent dusts and dose rates shown. 



Sample 



AdTL 
Code 



MiiZtn 


95042 


M02S/1 


<)\M*- 


MG2c/i2 


95044 


MG2<J/l2 


95045 


MG5S/0.I 


95039 


MC5S/03 


VMWili 




95040b 


M05S/1 3 


95041 


MG6S/60 


95046 


scios/U6 


95047 


SC12S/;. 


9VM8 



Analysis 

Healed 
Heated 
Heated 
Healed 
Healed 
Heau\i 

Hctiicd 

Healed 

Selective nleacb 
Sclceiin* hlgftCh 
Selective bleach 



Equivalent 
Dose (Gy) 



9.46 ±0.47 

9!3i0.32 

345*35 

82 ± IU 

15.7 i 1.3 

Mil 1 3 

23.9 ±2.5 

240 i 40 

14.t6±O60 

5,21*0.20 

2. 23 ±0.84 



Dose Rate 
(Gy ka 1 ) 



2 GO ± 0,03 

1.45 ±003 
1.39 ±0,04 
1. 50 ± 0.14. 
3.20±0.17 

3. 46 ±0.05 
3,46iO.05 
2.46 ±0.06 
1.74 ±0.06 
2/»3 ±0.08 
1.7*5 ±006 



Age (ka) 



3.64 ±0.25 

6.29*0*5 

250 ±30 

55 ±12 
4.91 ±0.4S 
4.08 ±0.38 
7.18 ±0.75 

98 ±15 
8.14 ±0.44 
1.58 ±0. 12 
1.27 ±0.48 



the quaru present jo this sample was actually in place 
well before the eruptions occurred. 

The 7 ka age found here is to be compared with that 
reported by Leaney et al. (1995) for a major change 
in the sedimcntology of the floor of the Blue Lake. 
However it is difficult ED reconcile the time scale ol 
lb** history of the Blue Lake a.s set by Leancy el <*/ 
widr ihe violent eruptive events traced by Sheard (1978, 
1990). One possibility is that Leaney er al have not 
sufficiently allowed for ihe incorporation of old carboa 
into their inorganic samples. 

The very large ages for the tuff MGIcIV *od 
MTOd/IZ collected from the cliff iace al the Blue Lake 
pump house suggest that this material d»d nnl in bci 
become sufficiently heated during the eruptions The 
very recent ages for the Mount Schank samples suggest 
thai there was some bleaching mechanism occurring 
1-2 ka ago. another date which has previously been 
associated with events at Mount Gambier, although it 
is now thought that the C-14 date on which this is based 
may he due in intrusive tree root charcoal (pers. comm. 
Blackburn to Sheard 1995), 

Because ol the problem in TL dating of determining 
whether the event that is being dated rcnuwed all the 
existing stored energy, all the dates should be carefully 
considered with regard to other Qvidence. For example, 
do the two apparent groups of ages obtained, i.e. e.4 
ka and c.7 ka really indicate two eruptions separated 
by 3 ka? During this time interval there should have 
been considerable weathering of any tuff ejected. 
However, the chemistry of the tuff sample.s MG5S/0.U 
MG5S/0.3 and MG2S/1 tuff (grouped together in Table 
2) shows that the ratio of the more soluble and mobile 
elements, sodium, magnesium and phosphorus to the 
insoluble element, titanium, is the same as that found 
for the solid Invn. Thus the lava must have protected 
the Riff very soon after it was deposited. Thctc are 
also no known palaeosols developed within this tuff 
or on top of the lava (Sheard 1990). The exposed tuff 
MG6S/60 does show considerable weathering, wilb the 
loss of about 50% of the soluble elements in relation 



to titanium. Incidcmly the Inst Of elements from 
MG6S/60 is about die same as the- loss of the same 
elements from the similar site SCtOS/O.b near Moiuu 
Schank. The age of the Mourn Sehank erupt inn was 
found to be 4.9*0.5 ka by Smith and Prcseotl (1987) 
using well baked tuff/sand under the lava How and ihe 
age of MG6S/60 should be similar. Those chemical 
considerations support the suggestion thai neither 
healing nor exposure to sunlight has been sufficient 
in sonic of the samples to remove all ol the prv-cxisttug 
TL 

M. y Sheard (pers. comm.) slates, "The phrcatic 
style of eruption so evident at Mount Gambier (i.e. 
associated wilh copious H,0) means that man)' 
eruptive products incorporating exotic quartz may nnl 
have been raised much above 100°C. ... In addition, 
under high volume teph.ra explosions some or most oJ 
the exotic quartz could escape light bleaching (i.e. ru 
setting) due to ash cloud or surge cloud density and 
subsequent rapid fallout and burial. Thus, it is possible 
to have exotic quail/ grains, only partially reset or left 
with their "older" TL signature, incorporated into much 
younger tephra" Sheard's remarks seem to indicate the 
possibility of finding samples containing partially 
heated quartz grains mixed witfi thoroughly bleached 
tephra. and so possibly being able to detect different 
dates tor difrefcm sued groin fractions. With this end 
in view, an attempt watt made to date the fine grain 
fraction of the samples from which qua/u grains had 
been extracted. Hut it was found in the ihree samples 
tested (MU2LVI0, MG2e/12 and MG2d/l2) that ciiheT 
there were insufficient line grains ol that the tine grains 
showed no TL and no sensitivity to radiation. 

The TL dates in conjunction with other geochrono 
logical evidence have reinforced the belief that there 
was volcanic activity at Mount Gambier at about 4 ka 
ago, and that then: may have been activity at 1.5 and 
7 ka. but that the latter dates should be accepted with 
caution. The iiivcstteation* have also illustrated tfut 



12 



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



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

Acknow ledgments 

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



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



References 



Aitkfn, M. J f (1985) "Thermolurninescence Dating" 

(Academic Press, London) 
BAKMtrn. M & Sheard, M. J. (1981) Palaeomagnetic 

results from Mounts Gambler and SchanK, South Australia 

/ Geol. Sac. £ Aust. 28. 385-394. 
Barton. C. E. & McEi.hinnf.y, M. W, (I9S0> Ages and 

ashes in lake floor sediment cores from Valley Lake, Mt 

Gambier, South Australia. Trans. R. Sac. S. Aust. 104, 

161-165, 
Blai krijrn, G., Allison, G. B. & Lkaniy. V, W. J. (1982) 

Further evidence on the age of tuff at Mt Cumbier, South 

Australia, Ibid. 106, 163-167. (With correction Ihid. 1984. 

108. 130). 
Ft;rgijsson, G. J, A RATOtt, T. A. (1957) New Zealand C M 

age measurements. N.Z. 1 Set. Tech. 38(b), 732-733. 
Hi NTirv, 0- J , HurmN, I. T & Prescott. J. R. (1993) 

The stranded beach-dune sequence of south-east South 

Australia: a test of rhermolunnnescenee dating 1 , 0-800 ka. 

QuuL Sri. Rrv. U, J ?0 
LhANhV, F. W, J,, AiitsoN, G. B., Dioiiton. J C & 

Tkumhork, S. (1995) The age and hydiologica) history of 

Blue Lake, .South Australia, Palaeogeog. , Pnfuct'tlim. , 

PuliwovioL 118. Ill 130, 
Nambi. K. S. V. Sl Aitkbn, M. .1 (1986) Annual dose 

conversion factors for TL and ESR dating. Archaeometry 

28. 202-205. 
Pktisroii. J. R.. Hunii-Ry, D. J. & Wtton, J T {19.93) 

Estimation of equivalent dose in ihermoluminescence dating 

- the Australian slide method. Ancient 77, 11. 1*5. 
& Hutton, .1 T (1994) Cosmic ray contributions to 

dose rates tor luminescence ami HSR dating: large depths 

and long-term time variations, Rtutiat, Mcas 2S. 497-500. 



& (1995) Environmental dose rates and 

radioactive disequilibrium from some Australian 
luminescence dating sites. Quat. Sri. Rev. iQuut GeOchwt) 
439-448. 

& Moiarkabl B. (1993) Selective bleach: an 

improved partial bleach technique for finding equivalent 

doses tor TL dating of quartz sediments. Ancient TL U, 

27-30. 
Robertson, G. B., Prfscott. J. R. <fc Hutton, J. T. (1996) 

Thermolnminescenee date lor lava flow at Mount Gambier. 

South Australia. Proc. Fifth Ausl Archaeometry Conf.. 

Armidale, 1994 in press. 
SimARD. M. J. (1978) Geological history of the Mount 

Gambier volcanic complex, southeast South Australia. 

Trans. R. Soc, Aust. 102. 125 139. 
(1983) Volcanoes pp. 7 14 In Tyler, M J.. Twidalc. 

C. R., Ling, J L,, & Holmes, J. W. (Eds) -Natural History 

of the South-East." (Royal Society Of South Australia, 

Adelaide). 

(1990) A guide to Quaternary volcanoes in the lower 

South cast of South Australia pp. 40 50 fn Drexel. J. E 
(Ed.) "Mines ami Energy Review South Australia." No. 157, 
(Department of Mines and Energy). 

(1995) Quaternary volcanic activity and volcanic- 



hazards pp. 264-26K fn Dtexel. 1F& Preiss. W. V (Eds) 

"The geology of South Australia, Vol. 2" South Australia 

Geological Survey Bulletin, 54_ 
Smith, B. W. & Prescott, J. R. (1987) Thermolurninescence 

dating of the eruption fit Mt Schank, South Australia. tost. 

.1 Earth Set. 34, 335 3-42 
Wintm-. A G & Hiintiiv H J (1982) Thermotummes- 

CCBCt dating of sediments. Quern Set. Rev. I, 31 53 



NEMATODES ASSOCIATED WITH DIPTERA IN 

SOUTH AUSTRALIA: A NEW SPECIES OF FERGUSOBIA 

CURRIE FROM A FERGUSONINID AND A NEW RECORD OF 

SYRPHONEMA LAUMOND & LYON FROM A SYRPHID 



ByKerrieA. Davies* & Janine Lloyd* 



Summary 

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

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



fhmmtWfnsttfthi'RomiSoaftvqfS. Awft. (1996). 120(1), 13-20. 

NEMATODES ASSOCIATED WITH D1PTERA IN SOUTH AUSTRALIA: A NEW 

SPECIES OF FERGUSOB1A CURRIE FROM A FERGUSON1NID AND A NEW 

RECORD OF SYRPHONEMA LAUMOND & LYON FROM A SYRPHJD 

by Kerrie A. Davihs* & Jamne Lloyd" 1 

Summary 

Davirs, K A- & Lloyd, J. (1996) Nematodes associated with Diptera in South Australia: a new .species of 
Fcrwxobta Currie from a fergusoninid and a new record of Sytplnmema Laumond & Lyon from a syrphid. Trans- 
It, Sm S. Ausr 120(1), 13-20. 31 May, 19%. 

f-ergusnhiti fishen sp. nov^ associated with a fly Fergusonim sp.. is described from leal galls on a hybrid 
of FMcaivptus U : ti(UK\hn. Like other species i)f Fit^usobia, these nematodes are semi-obese and have a generation 
parasitic in the lly followed by a generation parasitic' in the plant. The new species is distinguished from F. tutnifaek'ns 
by its smaller size, larger manuhrium on the spicule of the male, and smaller cephalic region ami tail m the 
partht-Tiugenetic female, with vulva and anus opening into culicular depressions, Syrpho/wnui sp., a nematode 
parasite of syrpnid flies, is described and recorded for the first time outside France 

Key Words; Taxonomy, Kematoda, Fcwsobia, Diptera. Syrpfwnema, new species new record. 



Introduction 

This paper describes a new species of the tylenchid 
nematode Fergusohut, the only genus of nematode 
known ro parasitize both a pJant and an insect 
(Maggenli 1981). It has a tripartite life cycle, with a 
generation parasitic in galls on Myrtaeeae followed by 
one parasitizing the fergusoninid fly Fcrgusonirui 
Mai luck: Also reported is. the first collection outside 
France of the rhabditid nematode genus, Syrpiuntttnu- 
Both nematodes were collected in Adelaide- South 
Australia. 

Materials and Methods 

Nematodes were collected from galls cut open in lap 
WatCT, relaxed and fixed in hot FA 4:1. Insect** were 
dissected in 0.85% saline, and nematodes from rhc>c 
were fixed as above. Nematodes were transferred from 
focative to \% glycerol in 30% ethanol m glass blocks 
and placed in a desiccator containing 9696 ethanol for 
2 days. For the following 14 days nematodes were kepi 
at 4(TC and ihe blocks were partially covered with 
glass lids During the first three days, one or two drop* 
of a solution of 5% glycerol in 95 ml cthanol were 
added four times daily. Slow evaporation was continued 
thereafter. For light microscopy, processed nematodes 
were mounted in glycerol on glass slides and examined 
using interference microscopy. Scanning eta Iron 
microscopy studies were made. Nematodes were taken 
from glycerol, passed through a series of 
ethanol /glycerol solutions with increasing proportions 



Department ol Crop Protection University of Adelaide 
PMH 1 Glen Osmond S. Aiist. 5nb4 



of ethanol and washed three times with 100% ethanol 
They were then dried using CO-, in a critical point 
drier, mounted on stubs, sputter coated with 
approximately 30 nm ol gold and viewed at 20 VV 
Measurements are in /ma. Drawings and measure- 
ments were made from material mounted in glycerol 
using a camera lucida. Body width was measured at 
mid-length. Spicules were measured along the mid 
line in lateral view. Dc Man's ratios, t.e V = anter 
tor end to vulva as percentage ol body length, T = 
length of testis trom cloaca to flexure as percentage 
of body length, a = length divided by gieatest body 
width, b* — length divided by distance from anterior 
end to base of oesophageal glands, e — length divided 
by tail length, c = tail length divided by width at anus 
wete determined. Comparisons were made with 
described species usinu published descriptions 
specimens of parthenogenctic females of Ferguselvo 
rfiiigtw Siddiqi (Queensland Museum (QM> 
G2U0512-20U51 4 *) and specimens of all stages except 
parasitic females of F Immtaaens Curne isolated by 
the authors i'tom flower bud galls on Eucalyptus 
cuxmald ulcus is Dehn. at Urrbrae, South Australia 
(Wnitc Institute Nematode Collection rWINO 943J 
The holotype of the new species is deposited in the 
South Australian Museum. Adelaide (SAMi 

Taxnnomtc descriptions 

fergusabia fisheri sp nov. 
(FIGS 1-2) 

tiolet\r[t€\ Parthenogenetic q. Black Forest, South 
Australia 1.34*57 S, |38°34'F.), 3vimr»3, W Frost. 
collected from a leaf gali ^\\ a hybrid of Euntlvfnu> 
facmylm E Muell., AHC207O5I (SAM). 



14 



K A, DAVTES & I LLOYD 



Paratopes: WTNC 815. slides [-28, including 20 
parthenogenetic 9 Q . 25 preparasilic infective 9 9, 
59 parasitic 9 9. 32 era, 59 juveniles, collected 
from leaf galls on a hybrid of E. leuoaylon at Black 
Forest between August and October 1993, by W. Frost, 

Measurements: Table I. 

Description of panhenogenetic female (Figs 1A, H, 
2A) 

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



part with valve-like structure, followed by short narrow 
"isthmus" which widens abruptly to broader part 
associated with the oesophageal glands; oesophago- 
intestinal junction obscure- Lumen of tract broadens 
posterior to "valve" and again at level of secretory- 
excretory pore. Oesophageal glands large, occupying 
about three quarters of body width, extending over 
intestine to about 55% of total body length; dorsal 
gland with large nucleus. Secretory-excretory pore 
50-92 fim from anterior end with short duct leading 
directly to excretory cell. Nerve ring at base of swollen 
anterior part of digestive tract, Hemizonid not seen. 
Reproductive tract with single gonad, prodelphic, 
extending to nerve ring; oviduct flexed in some 
females; uterus with quadricolumellar, often contains 
single egg, curves to join vagina, which is directed 
slightly forwards. Vulva usually conspicuous depressed 
transverse slit, but in some specimens, vulval lips 
protrude slightly. Cuticle wrinkled on ventral side just 
anterior to vulva in hall specimens examined. Rectum 
simple tube, without obvious musculature; anus 
inconspicuous pore, usually associated with distinct 
indentation in the cuticle. Tail conoid, narrowing 
sharply to rounded tip, 1.0-1.4 times as long as anal 
body width. Phasmids not seen. 



Table 1. Measurements of Fergusobia Fisheri sp, now 

Measurements in /tin, n for each measurement in brackets below mean 



tfolotypc Paratyp-es 

(parihenogenetic Parthenogcnetic females Males Pn:-parasitic females Parasitic females 

female) mean S.D. range mean S.D. range mean S.D. range mean S.D range 



Length 


250 


253 


23.2 


228- 


336 


9M 


292- 


349 


35.8 


241- 


786 


57.8 


690 






(12) 




305 


(16) 




453 


(25) 




395 


(20) 




905 


Width 


50 


37 
(12) 


2.1 


33-*0 


37 
(16) 


3.7 


32-46 


38 

(25) 


6.2 


28-49 


112 

(20) 


79 


105- 
128 


styict length 


10 


9.4 
(8) 


0.38 


9-iO 


y 
(13) 


1.0 


8-!l 


8 

(11) 


1.4 


6*11 


nil 


" 


- 


Anterior end to 


80 


75 


15.4 


50-92 


76 


12.2 


61-93 


75 


10.6 


6691 


- 


. 


, 


sec. -ex. pore 




(9) 






(6) 






(5) 












Vail length 


£ 


14 
(12) 




1316 


31 
(16) 




23-42 


30 
(25) 




21-49 


nil 


- 


- 


v 


87 


85 
(12) 


1.4 


83-88 








80 
(25) 


3.0 


76 88 


83 
(18) 


2.7 


78-89 


\ 


" 






■ 


75 
(16) 


7 2 


59-84 


-~ 


- 


• 




- 


- 


Spicule length 


- 


- 


- 


- 


18 

(17) 


18 


16-20 


- 


- 




- 


- 


- 


a 


5 


68 
(12) 


07 


6.0- 15 


9.2 

(16) 


1.4 


6.3-11.3 


95 

(25) 


1.7 


68-12.5 


7.0 
(20) 


0.6 


5.9 8.1 


b' 


1.8 


1.9 

(12) 


0.15. 


1.8-2.1 


2.8 
(10) 


0.41 


2.3-3.4 


3.2 

(9) 


0.42 


2.8-4.0 


- 


- 


- 


c 


15.6 


15.1 
(12) 


4.4 


1025 


12.9 

(16) 


1.5 


11-17 


12.9 

(25) 


4.4 


7-23 


- 


• 




c 


1.2 


1.2 


0,13 


0,9-13 






- 


- 


_ 


. 






_ 






(12) 

























NEMATODES ASSOCIATED WITH DIPTERA 



IS 



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

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



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




Fig. 1. Fergitsobia fishtri sp. nov. A. Entire parthenogentic female. B. Head of pre-parasilic female. C. Entire male (bursa 
nol seen due lo angle at which (ail viewed)- D. Detail 01 "male tail, ventre- lateral view, showing bursa and angulated spicule 
E. Entire parasitic female. F Entire pre-parasitic leinale. 0. H. I. Stylets of pre-parasitic female, parihenogenetic female 
and male, respectively. J, K. Lateral and dorsal view, respectively, of spicule. Scale bars = [0 p OHM K, 20 prn 
AKCD I*, 80 /mi E. 



16 



K. A, DAV1ES & J. LLOYD 




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

Description of parasitic female (Fig. IE) 

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

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

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



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



NEMATODES ASSOCIATED WITH DITTERA 



I" 



Diagnosis- anti relationships 

ft Jisfwri sp. nov. is characterized by having a 
parthcnogcnetk lernale with ibe cephalic region small 
relative to the body width, with a flat terminal pwfilr. 
the vulval slit in a conspicuous depression of ihc 
cuticle, ihc- anal opening in a similar but smaller 
depression and a short tail {0.9-] 3 times anal body 
width) with a narrow cone shape. The male has ait'tilai 
spicules with a large manubrium, 

F fishcri sp, nov. differs from F. jambophilu Siddiin 
in having a flat cephalic region without a central conk ;il 
elevation ami angulai spicules and front E itidtca 
(Jatrujpuri) and F. magna because the pjirthenogenetie 
female has a short tail, 't'hcauus opens into an obvious 
cuucular depicssion in the partficnogcnctie female of 
F fishcri sp nov bui not in the other described species. 
F fishcri sp. nov. assumes a similar shape '•> n* 
tumifiuifrts when heat killed but is smaller (avciagc 
length of parthenogcnctic females 253 and W jim 
respectively and ol males 336 and 42U fUM. 
Measurements for F, airriei ( = tumifacims) from 
Fisher and Niektc 196X>. The parthenogenctic female 
of F fishcri sp, nov. has a smaller cephalic region 
relative to body width tlvin F. tumifaciens and may have 
wrinkled cuticle on the ventral side of the body anterior 
to the vulva, absent in F. tumifiniens. The vulval slit 
is situated in a distinct cuucular depression in F fishen 
sp. nov. as m F. jumbupfalu and F tmltea, but not in 
F tumifaciens or in F ma$na. The volume of the tail 
ol the parthenogcnctic female is smaller in F fitheri 
sp. nov. than in F mtmfacienz. having a narrower cone 
shape The point of origin of the caudal alae is variable 
m E fishcri sp. nov. males, in contrast to F. nmufactem 

Ftymolo}>\ 

Named alter Vr J. M. FiSheT, formerly ol the 
Department of Crop Protection University of Adelaide, 

BtoloK\, life c\fft and general comment* 

The neinalodr F fishcri sp. nov. was found on leaf 
tfaJIs of the southern blue gum F. leacoxylotu in 
association tofth an unknown species of the 
fcignsonmid fly F$r#to$&litoa $p GftLuj were first 
(."Hot led hi cat ly August W93, followed by successive 
collections until October 199? when no nematode: |)I 
flics were found. In thcearJ) August collection, galls 
contained many nematodes (juvenile*, porthenopenetic 
and iiilceuve females and tuaios) and fly latvac Ol 
J3 Iflpfcfi tis>evted. the abdomen of two contains! ;i 
total of three fertilised infective females and one ittfljfc 
nematode. A week later, M tly lai vac and 15 piipaf la 
were collccied from calls and dissected. Su of the 
larvae contained two to four fertilized lnlevttve lemttlt 
nematodes. No nematodes were round in poptiriti, Jn 
early September, galls contained a lew adult male ami 
infective female nematodes. Puparia contained pbaraic 



adult flies. Nineteen pharate adult rites were dissected. 
eight of which were male and had no nematodes. 
However. 10 of the 11 female flies contained from I-ll 
(average 6.4) mature parasitic female nematodes per 
fly. In six of these, eggs had been laid in Hie 
Itaemolyurph and va most cases, the eggs were in the 
early stages of embryonic development. One fly. 
however, contained eggs in which the juvenile 
nematodes were well-formed. 

These observations on the life cycle generally agree 
with those reported by Fisher A Nickle <Wr>X) for F 
i vnii'i ( — tutmfiuiens) . No nematodes have been found 
in male flics. Eggs are laid in the hacmolymplt ol the 
abdomen ol the fly but ii is not known how the fl> 
deposits both insect eggs and nematode juveniles into 
Fucaty/nas tissue. Presumably, these nematode 
juveniles develop into panhenogenetic females which 
lay eggs It is possible that juveniles developing from 
the Batty eggs Inxome males, as in F rmnifaiietts 
While the first collection of F. fishcri sp. nov. from 
leaf galls yielded ali plant parasitic stages of the 
nematode, the fourth stage juveniles touud were all 
female (distinguished from males by a more bluntly 
rounded tail and development of the uterus) . suggesting 
that male development had occurred earlier. Fisher & 
Nickle (1968) stated that only fertilized infecrivestage 
females oft F curnti i^tttmifaa'ens) penetrated fly 
larvae but female and male F Jisfwri sp. nov. have been 
found in fly larvae. F rum t facie as may deposit eggs 
in the haemolympb of the fly before it emerges as an 
adult female (Currie 1937) or litis- may be delayed until 
after emergence of the fly from the gall f Fisher & 
Nickle 1968). Here, F fishcri sp. nov. had produced 
eggs before the parasitised fly had emerged from the 
puparium. 

Currie (1937) described one species, F. tumijuuern. 
associated with Fn. tarteri Tonn . from leaf galls on 
E, Stuariiana (sic) E v. M. He also observed minor 
morphological differences between ncmaiodcs 
collccied from leaf bud. a_\iJ bud, stem tip and flower 
bud galls associated with & number of other fly species 
on mote than a dozen species of cucalypt. He suggested 
4 'further work will show dun die nematodes assoc fated 
with the different flies have diHcrciiccs in structure 
whuh entitle them to be considered asddlet«nt ipexteV 
(turtle 1937). I hese differences included variation, ui 
the point of origin of the caudal alae and the size ffilq 
position of the oesophageal glands OtftorVatioi 
material held in the WINC confirm that Uk--- 
characters will be important in Sffeciefi diffrrvnl*:Ui;n 
Fisher & Nickle (19t>K) rcdescntvd h \ Wftm 
{ = iumi/at otn.\) froo> flower hud culls Oil r 
. unm)dnte'ft\i\ Debnh BT5SOCJfli*(l widi Fn uil. 
Tonn. Ot the other described speck-s m ftTguyebtote, 
Stddiqi I19S6) described A magna from FftralyptWi 
siem galls associarcxl with m unknown 11) best ami 
fmm soil under hmr trees. F iodica fumi soil in India 



K A OAVIES & J LLOYD 



and E jambophtla from flowei bud galls on Syzygiuw 
cumini (L. ) Skeets associated with En. syzy^ii Karris 

The Waite Institute Nematode Collection contains 
specimens of Fergusnbia collected from stem, bud and 
leaf galls on Eucalyptus from Adelaide and Mi 
Gambier, respectively, which appear to be undescribed 
species. Fisher (pers, comm ) round an undescTibed 
species of Fergusnbia in leaf galls on E, cumaldulensis 
associated with Fn. locklmrtt Tonn. Jt is apparent . given 
the different forms of galls, that each of these 
undescribed species of nematode is associated with % 
different species of Fcrgu&onirta . Bach, however, is not 
necessarily restricted to one species of Eucalyptus. 
Presumably the nematode has evolved mechanisms, 
which are probably host-specific, to escape the fly's 
immunological system during the generation in which 
it is an insect parasite Thus, Fergusnbia . like the genus 
Anguina Scopoli (Krall 1991) may have a high degree 
of fWt specificity, in this case, for the insect host. If 
so. Cunie (1937) and Fisher & Niekle (1968) may not 
have described the same species of nematode, given 
that they were associated with different species of 
Fergusnnina. Indeed, Currie refers to the margins of 
the caudal alae of males of his specimens of E 
tumifaciens as being slightly crenatc, a character not 
observed by Fisher & Niekle (1968) nor in specimens 
isolated from E . canuttdulensis by the present authors. 
Also, the oesophageal glands of the parthenogenetic 
female of the specimens from £. canuddulensis arc 
longer than in Curries description and drawing of F. 
nonifaaens. 

The structure of the digestive tract of Eergusobm 
remains unclear. Given the small sue of these 
nematodes, and their typical dark colouration, it is very 
drffictfll to distinguish the anterior parts of Ihc tract. 
Fisher & Niekle (1968) described the anterior part Of 
the oesophagus as swollen, narrowing abruptly 10 tbnn 
a short isthmus ai the level of the nerve ring, ihen 
hmadening again to contain the large gjands. They 
believed that the oesophago-intestinal junction Occurred 
31 about the level Of the secretory-excretory pore 
Siddufl (N80} interpreted the anterior swelling of the 
digestive tract as a "pseudo-pharynx" and believed ilntt 
the valve-like structure it contains marked the 



junction of the oesophagus and intestine. An electron 
microscope study of the anterior part of the digestive 
tract Of Fergusnbia is needed to decide which of these 
interpretations is correct 

Svrphnnerna sp, 
(FIG. 3(1 

Measurements: Table 2. 

Description of female fPig, 3C. D. b. K G) 

Nematodes straight or slightly C -shaped when 
relaxed by gentle heat. Cuticle has Longitudinal grooves 
with many fine transverse markings; often appears 
loose at head and/or tail; lateral lines large (31 ;<m wide. 
27 13 /im), with smooth ribbon like appearance and 
single central ridge, Lip region often indistinct; six 
separate lips, each with small papilla. Stoma cup- 
shaped: cheilosiom reduced; promesostom about 4 j/ru 
long, and about same width; thickening of cuticle on 
ventral side of metastom, which makes stoma 
asymmetrical, and may form a rhabdion, telostom 
present. Amphid openings at base of lateral lips. 
Oesophagus rectilinear; no true bulb, vestigial valve 
present- Nerve nng situated in posterior third of 
oesophagus. Secretory-excretory pore opening just 
behind nerve ring; prominent excretory cell just behind 
oesophago-intestinal junction. Deirid.s not seen. 
Hcmi?onJd just posterior to nerve ring. Intestinal lumen 
lined with refractive material from oesophago-intestinal 
junction to rectal valve; lumen wide in young females 
but narrower in older. Three rectal glands. 
Reproductive tract with single gonad, amphidelphic, 
with genital tube running anteriorly; ovoviviparous. 
long uterus, extending almost la pojnt of flexure of 
genual lube just behind oesophago intestinal junction, 
oviduct extending backdown dorsal side of body; small 
post uterine sac present; no true spermalheca. sperm 
not seen; vulva with transverse opening, well 
developed associated musculature; posterior vulval lip 
often more prominent than anterior; vagina not directed 
anteriorly or posteriorly Phasmids not seen. Tail 
conical, terminus variable (Pig. 3) 



T*BLt 2. Mcasutrmt m,\ jftf adult jrrrmtts .// syrphonema .*./>. isitfoifd jrom Mefaide. 

All measurement. 1 * in pril 





Length 


"Width 


Anterior 
end Ui haste 


Tfcll 

length 


Anrci t"i 
end to 


V 


a 


h 


c 








rf Mb 


HWi 


vulva 

BOO 










mean 


1 t/u 


44 7 


140. A 


*Ij6 


Z%i 


(06 


13 it 


n 


12 


I' 


10 


10 


12 


12 


12 


to 


10 


SO 


136 


M 


20.5 


164 


220 


5.7 


36 


1.7 


1 1 


rrr,;.' 


1 IHQfl 


n-s* 


jucni 


^ pii 


miM620 


fSf\9-8r\S 


25-36 


UH3.4 


IWitf 




NEMATODES ASSOCIATED WITH DIPTERA 

Description of juveniles (Fig. 3A, B) 



19 



Second stage juveniles 573 /*rn (473-624; n=8), 
third stage juveniles 830 ujii (806-873; n = 3); fourth 
stage juveniles 1062 fim (920-1260; n=7). 

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

Collector, host and locality 

The nematodes were dissected from (he intestine of 
two females of the syrphid fly Simosyrphus 
graiulirorinis (Macquart), collected on sow thistle. 
Sonchus L. sp. . at the Waite Campus of the University 
ol Adelaide, Glen Osmond in January and December 
1993 by Mr E. Soleyman Nematode specimens are 
held in the W1NC 687. 

Biology and general comments 

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

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



Fig. 3. Syrphonema sp. A. Entire fourth stage juvenile. B. 
Anterior of fourth stage juvenile. C Anterior of adult 
female. D. Entire adult female. E. Variable tail shapes ol 
adult females. F. Vulva and tail of female G, Stoma of adult 
female. Scale bars = 10 ^m B C E F G, 20 urn A, 50 /an D. 



20 



K. A. DAVIES & J. LLOYD 



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



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



Acknowledgments 

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



References 



Clrrie, & A. (1937) Galls on Eucalyptus trees. A new type 

of association between flies and nematodes. Proc. Unn. 

Soc. N.S.W. 62, 147-174. 
Fisher, J. M. & Nicklf., W. R. (1968) On the classification 

and life history of Fergusobia curriei (Sphaerulariidae: 

Nemaioda). Proc, Hefminihot. Soc. Wash. 35, 40-46. 
Krall. E. L. (1991) Wheat and grass nematodes: Anguina, 

Suhanguitui and related genera pp, 721-760 In Nickle, W. 

R. (Ed ) "Manual of agricultural nematology" (Marcel 

Dckker, New York). 
Laumond, C. & Lyon. J, P. (1971) Le parasitisme de 

Syrphonema intestinalis n.g., n.sp., aux depens des 



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

Maggenti, A. (1981) "General Nemalology" (Springer- 
Verlag, New York). 

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



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



ByE. M. Campbell*, C. R. Twidale*, /. T. HurroNf & J. R. Prescott± 



Summary 

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

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

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



TrunsuHiotts of the Hoyal Socu-ty ofS. Aust (1996). UUU). 21-36. 

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

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

Summary 

CAMPBtLl., L"» M . Twidaip.C. R., HurroN.l J.& pRKSccrrr. J. R. (1996) Preliminary investigations of dunes 
ot the Gawler Ranges province. South Australia Tmns. R. Sm: S, Aust. 120(1), 21-36. 31 May, 19%. 

Three lieltls of Junes have developed m Ihe recent past within (lit* Gawler Ranges in the and-semiarid interior 
ol South Australia, The dunes (lunettes, parabolic dunes, transverse Junes, linear dunes, climbing dunes and 
tolling dunc^j art essentially relic forms, were active ttbout 4000 years BP and are now stabilised by vegetation 
though strong winds still cause occasional sand movement. Some of the dunes demonstrate sand transport over 
distances of at least 25 km, The origin of Ihe various morphological dune types is discussed. Supply ot aahd- 
the moisture undent ol the substrate, the vegetation cover and wind speed and duvetton are oil important topography 
influences the morphology of the dunes in various ways and is fundamental to any explanation o! climbing and 
falling dunes. 

Key Wokos: Gawler Ranges, lunettes, parabolic dunes, transverse dunes, linear dunes, climbing dunes, tailing 
dunes, thermoluminescents dating, 



Introduction 

In the mid-latitude deserts extensive fields of sand 
dunes are restricted to plains. Sand dunes have, 
however, been reported from desert uplands where 
topographic obstacles deflect or funnel the regional 
airflow and produce deposilional forms and patterns 
different from the essentially regular and repeated 
formations found in the duneftelds of the adjacent 
plains (Wilson 1973; Smith J982). They include sand 
shadows ol various types, sand sheets, obstacle dunes 
and climbing and falling dunes (Planhol & Rognon 
1970; McKee 1979; Mainguet 1984; Greeley & Iverson 
1985). The Gawler Ranges, located in the arid-senuarid 
interior of South Australia, is a desen upland within 
which three fields of sand dunes have penetrated the 
valleys between the bornhardt massifs and in some 
areas have overridden the low domical Kills (Fig. la, b). 



Geologic Background 

The bornhardts of the Gawler Ranges are developed 
in a layered sequence of silicic volcanic rocks (mainly 
rhyolites, rhyodacites and dacites) of Mesoproterozotc 
age (1592 ± 2 Ma - Fanning et ai 1986). The volcanic 
rocks consist predominantly of subaerially erupted 
ignimbrites (nuces ardentes deposits), welded to 
varying degrees, and with local occurrences of basaltic 
lava and agglomerate- They were intruded by granites 
of the Hiltaba Suite (1485 ± 16 Ma - Creaser 1989 



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

t Deceased. 

* Department of Physics, University of Adelaide S AvM 
5005. 



cited by Blissctt et ai 1989; see also Flint 1993) which 
now occur extensively in the western part of the upland, 
in the Kondoolka and Hiltaba areas, as well as in small 
Isolated outcrops near Rokarha Homestead (H.S.) and 
Lake Everard H S. They ate also exposed to the W, 
SW and S of the Ranges. 

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



a u .'-'■.■■' rn i 



; ' ■:; :.■■■,: |, 



\ i-H !"* 



I.!. 4llKlli II*. 



' 



ist 



- j 



M . — - 

r 

_ - •• ' 

:■ i ■ 



WW 



>.!• I I ^ 

i. | i,l. ■ .. lBK»i 

r- >i mi 



ii _ 

* 




- ■•• 

,... "■'.,.. ■ , , 



j I 

\ 

I - ■ ... 



i 



-' 



., , i . . . 
* * 



* •'■,'! I 

'n ., hi. I| 



m,. _ 



Fig. la. The Gawler Ranges province, showing location of 
the study site in South Australia (inset), localities mentioned 
in the text and average annual isohyets tmm). The 
Corrobinnie Depression dutis.' samples analysed by Gostin 
were collected from dunes adjacent to the mad from 
Wirrulla to Hiltaba H.S. 



E M CAMPBELL, C R TWIDALE. J- T. HUTTON & I R. PRRSCOTT 



meridional sets, has been exploited by weathering and 
erosion to form the major valley systems of the Ranges 
(Campbell & Twidalc 1991). The summit and upper 
slopes of the bornhuidls are essentially devoid of any 
weathered materials, though isolated patches of regolith 



-1 "-■-;: 



■•■■■ 



li 

» " " ■- : 

i* ' "' Ik? 



•3£ C 

1 



: ! 



v 



rt £l 



B 

- i r J 












' ■ i ■ 






fig. lb The geology, including dunericlds, > if the 03w|er 
Ranges province (after William* 1994). A. Hieni Dunefiekl 
B Piccadilly Oimeltcld. C Moonaree Duneficld. D 
Beacon Ounefield. L, llkina DuncfieM R Scrubby Peak 
Dunefield 






Wr9. 







L IFH III 

I irfepi d'jnc* 

• '.VtjLi.U' ilurif 



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



'Campbell, E. M, (1990) Structure and surface in the 
Gawler Ranges. South Australia PhD thesis. University 



01 Adelaide (Uripub. ) 



are preserved on the lower slopes, as for example 7 
km east of Nonning H S . and in the valley floors drill 
cores and dam sections indicate that the volcanic 
material is weathered to depths Of up to 50 m. 
Individual hornbardts attain heights of over 460 m in 
the S, but they decline in elevation to the N (e.g. Nukey 
Bluff r 460 mj Mr Nott 430 ni. Bond Hill 336 m, 
Chitaniiga Hill 317 m, Nuckulla Hdl 256 m, Mortimer 
Hill 232 m - see Fig. la). The upland is bounded on 
its SW and S sides by an escarpment which, though 
dissected and rising only 150-200 m above the 
surrounding plain, is nevertheless, and viewed 
regionally, linear and imposing, so much so that the 
explorer E. J. Eyre was led to describe the Gawler 
Ranges as "a vast mountain mass rising abruptly oui 
of the low scrubby country" (Eyre 1845). To the N. 
W and H the relief amplitude diminishes and the hills 
peter out as the plains become more and more 
extensive. 

The bonthard(s of the Gawler Range province have 
evolved in two major stages. The first involved 
planution and fracture-controlled differential subsurface 
wealhering m Jurassic or earlier times, the second, the 
stripping of the regolith in the Early Cretaceous to 
expose bevelled domical forms. Remnants of the etch 
planation surface represented by the summit bevels are 
particularly well preserved in the south, There has been 
only minor erosion of the ranges since the Mesozoii. 
(Campbell 1990 1 ; Campbell & Twidaie 1991). 

Numerous salinas. including lakes Gairdner, Harris, 
Evcrard and Acrarnan and many small saline playas. 
occupy low lying areas in valleys and plains. The 
depression occupied by Lake Acrarnan is the site or 
an ancient meteorite impact (Williams 1994) The large 
lakes stand about 120 in above sea level, They are the 
termini of closed drainage basins mom of which are 
not much larger than the lakes themselves. 

Climate 

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

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



DUNES OF THE GAWLfcR RANGES 



23 



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

The Dunefields 

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

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

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

MOGNAHEL 



NONNING 



-n- 



Meait ninthly 

rainfall 



Jl 



1 



L. 



Mean nt> 
•a in uays 



J I MAM J .1 A SO NO 
Months 




j'f'm'a'm'.i.i'a's'oVi 
Months 



Fig. 2a. Mean monthly rainfall and mean number of rain days 
for Moonaree and Nonning (Bureau of Meteorology 1993a). 
Length of record: Moonaree - 108 years, Nonning - 90 
years. 





January 
N 



T 



duly 



1 . 2 o >20 km/hr 



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



WOO Wl ERA 



fk 



yfc 



January July 

CEDUNA 

A- yf 






January 
N 



July 



aa _30>30 km/hr 



1-10 



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



24 



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



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

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

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



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



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

The most southerly dunefield, the Ukina Dunefield, 
is part of the Kododo Dunefield of Smith (1976 2 ). 
Both the field and individual dunes trend NW to SE 
between the Corrobinnie Depression (Bourne et al. 
1974; Binks & Hooper 1984) and the SW margin of 
the Gawler Ranges from near Yarranna Hill to the 
vicinity of Mt Sturt. Within the Corrobinnie 
Depression, complex parabolic forms are well 
developed. In the vicinity of Mt Centre, linear sand 
ridges from the Ilkina Dunefield diverge ESE and 
extend across narrow plains and valleys between the 
volcanic uplands and extend into the hilly areas to form 
the Scrubby Peak Dunefield (Fig. lb). In both the N 
and S arms of this dunefield there are departures from 
the general ESE trend as a result of topographic 
interference with the airflow. Both crestal transverse 
dunes and climbing and falling dunes result from such 
topographic effects. 

Dune morphology 

Linear dunes 

Linear sand ridges dominate the dunefields within 
the Gawler Ranges (Fig. 3). These linear forms trend 
WNW to ESE in the W and latitudinally further to the 




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



DUNES Or THE GAWLER RANGfcS 



15 



E. In places, e.g. near Ml Granite (Fig, 4). funnelling 
of the wind has produced dunes aligned at various 
angles m the regional trend. The linear dunes vary in 
height, length and linear frequency, i e, the number 
of sand ridges per unit distance measured normal 10 
the dune trend. The maximum height of the dunes 
varies from 5 15 m above the intcrdune corridors. They 
vary in length froW a few tens of metres up to 20 km, 
none extends unbroken for many scores or hundreds 
nl kilometres as do some of the sand ndges of desert 
plains such as the Simpson Desert (AVopfner & TwidaJe 
No 7, W90; Twidale 198!). The linear frequency of the 
dunes varies between two and six per km The 
intcrdune corridors *ire sand covered. Mos( Oft the dune;, 
are symmetrical, with smooth crests which rise and 
fall to form peaks and saddles. The slopes are gentle, 
considerably less than the angle of repose of the sand. 
No depositioual structures and no slip faces have been 
noted. The dunes carry a covering of low shrubs and 
small ttees. though little or no soil development is 
apparent and there is. today, only occasional and minor 
reworking of the sand by wind and water. The dunes 
arc relic according to the classification of Livingstone 
& Thomas (1993). 

Parabolic dunes 

Groups of parabolic or U-dunes occur within the 
linear duncfields. Most of the parabolic dunes occur 
outside the Ranges, and notably in the Corrobinme 
Depression (Bourne et ai 1974), though mere is a W- 
E zone within the Moonaree Dunefield and patches 



-& 



I Ffav i- 
_■ [Jmw 

l(HJ r".( IlluUf 



■■ ' , 
1 : > I 

'' \ . 



Fig. 4 Schematic diagram of linear dunes oriented NW ui 
SE, and irregular patterns of linear and transverse dunes 
due to topographic interference lo the wind ncur Ml Gi anile, 
Gawler Ranges, South Australia (from aerial photographs, 
Department of Lands, South Australia and H00 000 
National topographic map series). Not all dunes are shown 



of parabolic forms occur within the Piccadilly and 
Beacon dunefields. Although many of these dunes are 
complex in plan form, with transverse, rake like and 
circular patterns well developed, the basic unit is it V- 
shaped dune about 8 m high and with the open end 
of the U pointing, to the W (Hg, 5)- 

Cltmbing and falling dum<> 

In the Gawler Ranges most of the dunes gig 
developed on broad valley floors between the 
bornhardts. In some areas, however, linear dunes 
penetrate into the hilly terrain and suffer modification 
as a result of funnelling and diversion of the wind (Fig. 
4)- In other areas, the dunes extend over the bombards 
On the reasonable assumption that the sand migrated 
southeasiwards, dunes piled against the windward 
(northwestern) slope of a hi 1 ! are termed climbing, or 
rising, dunes; where sand has overridden the crest ot 
a hill and extended on to the leeward (southeastern) 
slope, tailing, or hanging, dunes are formed (Fig. 6a). 
v 




Fig. 5. Linear and parabolic dunes qf the Moonarec Dunefield 
15 km south of Lake fivtrard H.S, (from aerial photographs 
Department of Lands, South Australia and 1100 000 
Nabonal topographic map scries), 



• 



-.: -" 



_ 



:- 



Kig. 6a. Echo, climbing and falling dunes (aftei Mabburt 
1977). Arrow indicates direction of the wind A. Linear 
dune not anchored by topography. B. Linear dune rising 
over topographic obstacle. C, Climbing (1) and (ailing (2) 
dune. D. Echo dune. 



'(, 



n M. CAMPBELL, V H TWtDALC. J. T HUTTON & J R PRESCOTT 



These dunes have not been studied in detail in (he and 
mountains of Australia, although dunes which arguably 
ascend cliffs have been studied in the coasul context 
(Jennings 1957: Langford-Smilh & Thorn 1969) 

Climbing and falling dunes are known from various 
parts of the world, for example from pengiaeial Finnish 
Lapland (Seppala 1993). coastal west Galicia. Spain 
and NL Spam (Cros & Sena 199.3 > . but most 
previously published reports pertain to warm desert 
environments, e.g. California (Evans 1962; Smith 1967 
cited by Bender 1982. Anders 1974 cited by Bender 
19X2; Lancaster 1994). Colorado (Johnson 1968), 
Idaho (Ko.scielniak 1973"), Arizona (Giecly & JveTvo 
1985) and Utah (Xlhbrandt 1979), all in the United 
-Slates, where most are inactive, veneered by gravel and 
dissected by ephemeral streams (Smith 1982), northern 
Mexico (Stone 1967). Brazil (Bigarella 1975, rW), 
Egypt and Jordan (MeKee 1979K the Sinai Penintmly 
(Ahibrandt 1979). die Sahara (Smith 1954) and the 
central Namib Desert (Goudte 1972). They arc also 
found in the eastern Flinders Ranges, South Australia 
(Green 1994 4 ), near Port Stephens and in the 
Shoalha^en Rivet area in New South Wales (Thorn ft 
u! 1994), on Uie Erldunda Range 160 km south of Alice 
Springs and on the northern margin of the Simpson 
Desert where dunes override some of the latitudinal 
ranges. Greeley (1985. Fig. 7-39) illilstratct a field ot 
climbing dunes drifting aver the rim of a 16 km 
diameter crater on Mars. 

In the Gawler Ranges climbing and falling dunes 
occur in three areas. First, examples were noted by 
Smith in the Scrubby Peak Dunefield (1976 2 ; Fig. 6b, 
c). Second, Giles (1980 5 ) remarked that sand dunes 
encuKich on to the slopes of Mt Stuxt. Sand from the 
llkma Dunefield has accumulated on the NW slopes 
of Mt Stun (the western peak) and forms an irregular 
mound along the base on its SE side. Third, climbing 
and falling dunes are Common in the Moonaree 
Dunefield E of Lafce Atfaman, where small dunes 
trending W-E are essentially restricted to the plains, 
though they partially override many of the bornhardts 
(Fig. 6d)_ 

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



3 Ku.scju.niak, D. E. (J973) Eolian deposits on a volcanic 

terrain nearSaini Andumy, Idaho. MA thesis, University 

of New York (Unpuh.) 
Grpfwk, S. J. (1994) A geomorphological and sedimento- 

logical study of a climbing dune, northern Flintier* Ranges, 

South Australia BA (Honsl thesis, University of Adelaide 

(L'npub.) 
- s Gtixs. C W (19H0) Spring Hill, southern Gawler Ranges. 

Geo! Sot Aust. S A- Div Geological Monuments 111, Pile 

E 20 tUBDiih) 



classed as climbing dunes. In some instances the dune 
is diverted around the flanks of the hill and continues 
downwind (Fig. 6b) Elsewhere, the dune form is 
interrupted, tor although ihere are many grains and 
even small pockets of sand in fissures and shallow rock 
basins on the crests and upper slopes of the hills, there 
is no dune form; a short distance downsJope from the 
crest, however, the dune form is resumed m tailing 
dunes 3-4 m high (Fig. 6c>. 

Trtmsverse dunes 

In die Scrubby Peak Dunefield funnelling uf the wind 
has produced dunes of varied orientation. In this pait 
of the Gawler Ranges elongate bornhardts are aligned 
essentially N-S. Sand ridges also aligned N-S arc 
trusted in the valleys between the bornhardts. There 
are some W-fl dunes which override the bornhardtv 
and, in addition. N-S trending elongate dunes are 
located just below the crest on the lee side of these 
hills (Figs 4, 7), These crcstal dunes are tentative! 1 , 
classified as of transverse type 

Lunettes 

Lunettes are developed along at least part of the L 
side of most of the large sahnas and many of the smaller 
ptayas in the region (Fig. Ic). Lunettes arc transverse 
dunes located on the lee shores of lake basins. The 
name "lunette" was first applied to the form by Hills 




'"> - :■ i 



1* ",..;,.! -,,. 



i alltn j 





W 




Fig. 6b, Scrubby Peak area, Gawler Ranges, South Australia, 
showing diverted dune (D). climbing dune (C) and falling 
dune t_f) (from I; 100 OCX) National topographic map series) 
The ■dune-forming winds were from the northwest sector 



dunes of the: gawler ranges 



27 




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




■■■■■.:■■:■ : : "v 



smSmmWmm 



:;v;:-.r ■;'.rv.:;)- ; :::;r-r---' i '~ : : i 




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



.'K 



E. M. CAMPBhXL, C R TWIPALF, J. T. BUTTON & 1 R PRESCOTT 



(1940) who described lunettes of silty-clay 

compositions IVorn NW Victoria- Subsequent! v, 
lunettes of various sizes and mineralogies have been 
reported trom all states ol" Australia They ratine in 
composition from quartz-rich to clay-rich to almost 
pure gypsum The sandy quartz-neb dunes were 
formed hy deflation from beaches on the lake maxgrn. 
The clay-rich dunes were derived by deflation of clay 
a^rcgatcs from ihe saline lake floors The gypsum 
dunes are composed eiliter of rounded crystals deflated 
from the dry lake bed or of tine 'kopL some of which 
may be due to Weathering of saJlated particles since 
deposition (Stephens & Crocker l'>46; Campbell 1968, 
BoWfer 1%8. W83; Chen e/ ai 199M. 

In Ihe Gawler Ranges area lunenes of gypseous 
composition occur on the eastern margin of lakes 
Kvetard, Hatns, Acraman and off manv of the smaller 
Salinas. Both gypseous and siliceous lunettes are found 
on Ihe eastern side of Lake Gairdner. The mos! 
prominent siliceous lunettes arc located opposite the 
dunerlclds which impinge on the W side o\' the lake_ 
Both duuefields found G of Lake Gairdner, the 
Piecadilly and the Beacon, are developed in Ihe lee 
of these pronnneni siliceous lunettes. The lunette on 
the NE margin of Lake Gairdner nses about 35 m 
above the lake bed- Much of the surface is bare and 



erosion by wind and water has created a series ol 
domical remnants, standing 3-4 m above gentle swales. 
In addition, lunettes consisting predominantly ui 
fiagments of Gawler Range Volcanics of sand stye 
occur discontinously along the margin of Lake 
Gairdner (Fig lc). 

Sedwientology 

A total of 16 sand samples, each from the crest ol 
a dune, and including at least one fmm each of the 
duuefields in the Gawler Ranges province, was 
examined to determine composition and grain 
morphology vTahie I) <tv\i$ grain size and related 
parameters Wert* determined using 0,5 phi standard 
sieves. 

The sand samples are all various shades ot yellow- 
red (2.5 to 10 YR Munsell Soil Colours). All sample* 
consist of at least 90%. and most more than 98%< 
quartz grains. The minor constituents are quartz rock 
feldspar, Gawler Range Volcanics fiaginetus. ?irot) 
oxide and organic matter, hi most samples- the grains 
arc predominantly frosted, but some are polished 
Samples from two dunes in the Scrubby tfeak DunefielU 
show higher percentages of polished grains. Grains 
from all samples show ferruginous coatings of yellow 




%. 7 TVynsven-e ercstaJ -June. Scruhhy Weak Dunettclil. Gawlei Rank's. South Australia. View to the >uuih |u ih<* t~Y'rr<n.mm 
L'epresNioo. The crcstil dune la about 8 m high 



DUNES OF THE GAWLER RANGES 



29 



Tabli 


1 . Composition and gram morphology of sample. 


from dunefields in the Gaw 


ier Ranges, South Australia. 


Sample 


Dune type 


Colour 1 


Composition 2 


Surface 
texture 3 


Surface 
coaling 4 


Roundness^ 


Sphericity 


Beacon Dunefield 


2.5YR 5/6 


quartz 95% 


85% MF 


Y, O, R, B. 


SR-SA. 




1 


linear 


high. 








(1,2,3,4.5) 


15% P 




tew R 


some elongate 


Piccadilly Dunefield 














2 


linear 


SYR 4/6 


quartz p93 


98% MF 


Y. O 


SASR. 


high, 








(1.2,4,5) 


few P 




lew R 


some elongate 


3 


parabolic- 


SYR 5/6 


quart? 99% 


98% MF 


Y, O 


SA-SR. 


high. 








(4,5) 


few P 




few R, WR 


some elongate 


4 


linear 


2.5YR 4/8 


quartz 99 % 


95% MF 


Y, O, R 


SA-SR, 


high. 








(4,5) 


few LF 
tew P 




few R 


some elongate 


5 


linear 


2.5YR 4/8 


quartz 98% 


98 & MF 


Y. R, B 


SA-SR. 


high. 








2% P 




few R 


some elongate 


liiern 


Dunefield 














6 


linear 


5YR 5/6 


quuru 99% 


98% F 


Y, O 


SA-SR. 


high. 








a.4.5) 


2% P 




tew WR 


some elongate 


7 


irregular 


5YR 5/6 


quartz 99* 


95% F 


Y. O 


SA-SR. 


high. 








(4,5) 


5% P 




few R 


some elongate 


8 


linear 


7.5YR 6/6 


quart/ 99% 


95% MF 


Y, C) 


SA-SR. 


high. 








(1.4,5) 


5%P 




few R, WR 


some elongate 


9 


linear 


5YR 5/8 


quartz 90% 


90% F 


Y. O 


A-SR 


moderate. 








(1.3.4.5) 


10% P 




some R 


some high 


Moouaree Dunefield 














10 


linear 


7.5YR 5/6 


quartz 98% 


90% KF 


Y, O 


SASR. 


high. 








0,4,5) 


10% P 




some A. R 


some elongate 


11 


irregular 


7.5YR 5/6 


quartz 99% 


95% F 


Y, O 


SA-R, 


high. 








(4.5) 


5% P 




few WR 


some elongate 


Scrubby Peak Dunefield 














12 


linear 


7.5YR 5/6 


quart/ 95% 


95% MF 


Y, R t B 


SA-SR. 


high. 








0,4,5) 


5% P 




few R. WR 


some elongate 


13 


linear 


10 YR 5/4 


quartz 99% 


70% MP 


Y, O, H 


SA-SR, 


high-moderate. 








11.4,5) 


30% SF 




some R 


some elongate 


14 


linear 


10YR 6/4 


quartz 99% 


50% P 


Y O 


SASR. 


high-moderate. 








(1.4.5) 


50% F 




few R 


some elongate 


1? 


linear 


7 5YR 4/6 


quartz 98% 


98% F 


Y, 


SASR. 


high-moderate. 








(1.4,5) 


2% P 




few A. R 


some elongate 


Ilkina 


Dunefield 


7.5YR 6/6 


quartz 98% 


70% LF 


Y O, R 


SA-SR. 




16 


linear 


moderate high. 








(1,3,4) 


30% P 




few R 


some elongate 


'Munsell Soil colours. 














-Minor constituents in brackets: 1: quart 


i rock 2: feldspa 


r 3: Gawler Ra 


nge Volcanic* 


4: '.'iron oxide. 5: 


organic material. 



'F. frosted. P. polished. M: moderately. L. lightly. 

*Y: yellow. O: orange. R: red. B: brown. 

3 SA: subancular. SR: subrounded. A; angular R: rounded. WR; well rounded 



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

The dune samples are all fine to medium grained 
sands (mainly 0.125 to 0.5 mm diameter - Polk l%8). 
They are well-sorted to poorly sorted, with most 
samples moderately well-sorted. 



Age of the dunes 

The sand of the Gawler Ranges dunes is typically 
a yellow-red colour (Table I ) , suggesting sufficient time 
tor initial weathering of clay, release of iron, and 
development of a taint ferruginous patina. The sand 
is not the brilliant red of the deserts of central Australia, 
nor the dusky red (I0R 3/4 Munsell Soil Colour) of 
the sand derived from the local Gawler Range 
Volcanics. Some authors would attribute the contrasting 



M) 



U. M. CAMPBELL, C R. TWIDALU, J. T. HUTTON & 3 R. PRESCOTT 



colour lo different source materials (Wasson 1983; 
Nanson eta!. 1992). Others, e.g. Wopther & Twidale 
1 19&7) and Walker (1979) consider that the intensity of 
the red colour increases with lime and hence is an 
indication of the age of the dunc- 

In an attempt to attain a more precise estimate of 
age, sand from the Scrubby Peak Punefield was tested 
lor thermolaminescence (TL). Samples were taken 
from a tailing dune on an unnamed hill (National 
Topographic Map Series Minnipa 5932, 1:100 000, 
Grid Reference N[£3I6018) 1.5 km **: of Scrubby Peak 
(Tig. 6bL The method was. a variation on the "partial 
bleach" method developed tor ihc TL dating of 
sediments by Wintle and Huntley (1982) The age is 
estimated by measuring the TL energy stored in the 
lattice of a suitable mineral, in this case, quart/. The 
time interval measured is the time since the Stored 
t-ri^y was last reset 10 zero or near zero by exposure 
to Solar ultra-violet radiation After such a re-setting. 
energy accumulates again at a known rate by exposure 
to radiation in the environment from the naturally 
radioactive elements K, U. Th and from cosmic rays 
The age is found from the so-called age equation: 

iigc = natural TL 

TL ptr unit dose x dose rate. 

Samples were recovered from depths of 35 and 70 
cm within the dune by means of an auger, taking care 
to shield the sample from light during and after 
collection. After digestion with 20% HC1 to remove 
carbonates and NaOH to remove clay, the 90-125 /mi 
Iraetum was rccovcrvl by sieving. A 40 minute etch With 
40% HP removed feldspars and a suriace layer of the 
quail/. Rotation on aqueous sodium polytungstute at 
a relative density of 2 67 followed; the end product 
was very pure quart?; and it is on this sample that the 
measurement-* were carried nut 

One of lite problems with TL dating nl se-diimmi.. 
is uncertainly nt«».it the degree ro which the TL was 
icsct at the beginning or rhc rime of interna Ii is r.m- 
for die TL to be removed L-ompletclv. even fcj 
prolonged exposun. tQ sunlight. Moreover, the amount 
Dd ictic TL vaa *cs In itn sample io sample and may vary 
with the age of the sample ( Bcrger 1990), In the present 
u^oOgatioti, it was luund lhat the accumulated TL 
was small so Hut any uncertainty in ihe degree of 
resetting v/mdd result m significant uncertainty in die 
i,ec The level ot rtneiimg was round from a surface 
vamplc collected by pressing packmc tarn- against die 
exposed dune surface TtilS flawed that theTL clock 
had not been completely reset to zero in spite of the 
long litlte likely |0 have been spent in Ihc sun by rhc 
sample in reaching its present position. Under tliese 
•-iicumsUnces special procedures aic «ec>ii» . 
uWrihed by JVcst'ott and Mojarrutw tiM93i They u$akt; 



use of the fact that many quartz samples have a iff*- 
called Vapidly bleaching" peak (RBP) at 325 °C in the 
thermolumweseence glow curves, which bleaches to 
near zero within a matter of minutes when exposed 
to light of wavelength longer than about 500 nju 
(Spooner et at- 1988). This means that exposures ol 
the order of minutes to natural sunlight in thr. 
environment will have ensured that the trap concerned 
had been emptied completely and that, a,t least so tar 
as the 325 °C peal £ concerned, the TL clock of the 
sediment was completely reset at the lime of 
deposition. Jn addition this peak emits in a wavelength 
hand centred near 420 nm. so that an optical filter 
transmitting this band will he selective for the peak 
in question (Preseou & Fox W0; Scholeficld et at 
1994), The 325 "C peak rides on an unbleached 
background, which is measured and allowed for by die 
procedures The suriace sample mentioned above haa 
zero TL when measured with the revised proceduces. 

the TL is expressed in terms of an equivalent Jos. 
measured ingrays (Gy). The equivalent doses art: (ui 
the 35 cm sample 1.24 ± 0.20 Gy; and for Ihe 70 cm 
sample 1.53 =t 0.25 Gy. The dose rate has been 
measured by three essentially independent methods 
(Hut ton & Prescntt 1992). They are. with the relevant 
dose tates in brackets: in situ gamma ray spectrometry 
(0.153 ± 0.028 Gy ka '>; thick source alpha counting 
for U and Th with X-ray spectrometry (XRS) for K 
(0.142 ± 0,029 Gy ka 1 ); and delayed neutron analysis 
(DNA) for U. neutron activation analysis (NAA) for 
Th with XRS for K (0.165 ± 0.041 Gy fcr'i Tbe 
weighted average is 0.152 * 0.010 Gy ka l for both ihe 
35 cm and 70 cm sampler. 

Contributions lor commie ray> must be included. 
These art- 0.21 i 0.02 and 0.18 -t 0.02 Gy ka 1 for 
the 35 and 70 cm samples respectively (Preseott A 
Hutton 19X8, l UQ 4L |r is worth noting that cosmic rays 
dominate the dose rate because the levels of K, V and 
Th an: $g nuicmdy low (R - 0.04 £ 0.01T-; U 22 
t 0,0f> ppn>, Tl» ■ ID t 0.4 ppm>. Over the time in 
question, changes in cosmic ray mtcasilit:,-, are 
negligible (Prescott & Hutton 1994*. 

The dose rates are 0.35 ± 0.03 Gy fcu' at 35 em 
and 0.3-3 ± 003 Gy ka ' at 70 Cm, A contribution 
Fpftm systematic errors- ha^ oeen added. Hence ihc -ic<.- 
Of the 35 cm sample »s 37 ± 07 ka and of the 7(1 ;m 
sample 4.6 ± 0.9 ka. Although the deeper sample has 
the- greater TL age, the IV Q age* arc nor statistically 
different urni probably all thai can he concluded is ih.v 
die dune has been in place for about 4 ka 

This age is based on 8 sfc^gfe series ot date* [mm 
one dune Obviously, more age doiemunanons aire 
required. Nevertheless, the pale colour of the sand, 
to which previous reference has been made, mid the 
lack ol Any ..trhormte -iccomulaltons Lfl llic duties 
,• itc itsiAailabiliryJa/esUggpMiveofayoulhful age 



DUNES OI : THli G*WU<R RANGES 



31 



The general appearances arc consistent with the TL 
dating in suggesting thai the Gawler Kanges tilings jre 
younger than the putative Late Pleistocene relic ,>rms 
of NW Eyre Peninsula (Twulale cr at, l*J76> and are 
comparable to the Holocene forms of (hat area (Rankin 
& Flint 1991) and of the Simpson Desert {Wbpfner & 
Iwidale 1988, 1990). Nevertheless, the age 
determination obtained is for the uppermost Ia>vr-. of 
a dune and there is no evidence of the age nf the sattd 
at the base of the dune. 



Origin of the dune sand 

As mentioned previously, the provenance of the sand 
in duncficlds. whether it is of local derivation or far- 
travelled, is controversial. The question can be clarified 
by a consideration of the sedtmentologic characteristics 
of the dune and otheT sands. In the Gawler Ranges 
province, a local origin of the dune sands is precluded 
by their composition and granulometry as set out in 
fable I For example, the Scrubby Peak Dunefield 
overlies outcrops of Eucarro Dacite, and Yardea Dacite. 
with small areas of Yannabie Rhyodacite, Pancy 
Rhyolite and, at the base ot the Yardea Dacite, ''black 1 
dacite (Blissett er ttL 1988, Fig 8)_ The microscopic 
groundmass of the volcanic r^ks is rich in quart/., but 
the grains are much smaller than rhosc of the dune 
sands. There are no quartz phenocrysis in the dacite 
(Blissett 1986). No quart/ of a size equal to, or greater 
than, thai of which the Scrubby Peak dunes are 
composed (and hence susceptible to attrition to produce 
sand- sized grains) could be derived from the Yardea 
and Eucarro dacites which .ire the country rocfc over 
winch, overwhelmingly, the duncficlds have extended 
No lakes or streams which might constitute a possible 
source of sand in the dunes are known from within 
the province. 

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

First, there are outcrops in 'the western Gawler 
Ranges of Yannabie Rhyodacite and of Paney RhyoJile 
(Fig. S), both of which contain pheuocrysts of quartz 
of a size equal Id, or greater than, the dune sand (0.2 
to 2.0 mm in diameter - Blissett (98b) Similarly, and 
second, granite with abundant coarse quartz crystals 
crops out to the west of the Ranges (Blissett <>f ai 
1988). But difficulties attach to these outcrops us 
soutces of the dune sand: they arc of limited extent 
(about 60 km 2 compared with the 300 km 3 of the 
Scrubby Peak Dunefield); and it can be questioned 
Whether they could produce a volume of quartz sand 
compatible with that represented by the total of the 
dunes. Also the outcrops do not extend across the width 
of the dunefield . so that the spread of sand from them 



calls for varied strong winds, and lor distribution in 
topographically difficult terrain Moreover, the 
dunefield extends westwards, i.e. windwards, of the 
outcrops in queslion (Fig. 8). Against these arguments, 
the Gawler Range Volcanics form a regional basin 
structure so that before erosion to their present 
occurrences, the quartz -bearing members could have 
extended further to the W. In addition, the former shape 
and size of these members could have been very 
different from their piesent representatives. But. on 
the evidence, the rhyodacite. rhyolite and granite 
outcrops Of the western ranges and adjacent areas do 
not seem likely sources <y. suitable quartz sand. 

The third possibility is that the dune sand has been 
derived from the Corrobinnie Depression, This runs 
in a NW-SE direction wes>, and therefore windward. 
of the dunclleld and contains detritus derived from the 
granite areas to the S. W and N as well as trorn the 
Gawler Ranges. It contains quart? comparable in size 
and character to the dune sand of the Scrubby Peak 
Dunefield (fine-grained, moderately well sorted, 
typically subrounded, frosted and coated with iron 
oxide - Gostin pers. comm 1993). it is concluded that 
the dune sands of the Scrubby Peak Dunefield cannot 
have been derived from the disintegration of the Gawler 
Range Volcanics. but rather have been transported on 
the wind from the Corrobinnie Deptess'on- a distance 
of at least 30 km. Even if derived from the Thyolite, 
rhyodacite and granitic outcrops, the sand must ha\e 
travelled 25 km to cross the zone of dactttc bedrock 
Given the wind regime, the northern arm of the 
Scrubby Peak Dunefield (\ in Fig 8) could only have 



i \ 






', A 




• 


- 




. 


■ 


- s (ii 


,i. 


"1 


.Tl 


• 


\ 


X 


> 


* 1 


J» 




\ 




! 


, , Hi 


A 


I 


| r. . .■■ ■!, 

1 . . | . i ... 






1 AM .i)f| 


•^ ~~~*^\ ■ *' ' 


i . , . . .1 




" k ^— — — . 1 



Fig. 8. Scrubby Peak Duneield, western Gawler Ranges, 
South Australia, illustrating bedrock type and possible 
sources of the June >.*m»t X. Northern ami of ihe 
Dunefield. Y Nearest upwind outcrop o( rhyoliie/rhyn- 
daerte. ED. tiucarro Dacite YD- Yardea Dacite R. 
Rhvtjbit: RD RhvoitxjR' Sec tcKt for explanation. After 
Blissett W ui 19KX 



32 



I V t-AMPBELL. C. R TWUMl.E. J T H'.'ITON & J R PRESCOTT 



originated in (he rhyolitic/gramtie outcrops indicated 
by Y in Fig. 8 or from outcrops further wv.siw.in1 
Whether this would he considered far-travelled l& -a 
matter of definition, but the sand is certainly not of" 
local denvution. 

Median ism of dune formation 

Prevailing winds 

h is suggested (see bctow) that the duneJVIds of (ho 
CJawler Ranges have been shaped by wini* from the 
western sector. TTO is consistent with llie putative 
source of the sand of which the dunes :ire constructed 
(see below). The relic linear dunes to The south of the 
Gawler Ranges, which possibly formed in late 
Pleistocene times, extended from NW coSE across the 
northern base of Eyre Peninsula, for they extend on 
to the svestern shores of salina.s such as Lake Agars, 
but not on to the eastern shores (Twidale & Campbell 
1985). These NW-SE dunes also extend well below 
low tide level between Co well and Whyulla (Van Peur 
1983*) but only in minor degree on the eastern sidt 
of the Gulf on northwestern \brke Peninsula, where 
the aeolian forms were deposited during a phase of 
rising sea level and where the dunes 3re truncated by 
wave action at the coast (Jessup 1967. 1968). This is 
consistent with a wind regime dominated by north- 
westeTlies. 

In addition, at Lake Gairdner, the lunettes of the 
eastern shore are much more substantial than those of 
the western and, as the lunettes are comparable Do 
coastal foredunes (Campbell 1%8), this supports a 
westerly wind regime. Also, die tiuge Late Pleistocene 
calcareous aeolianite riiredunes of wcsn-facirig shores 
in Soudi Australia (e.g. on Eyre Peninsula) dwarf their 
east coast counterparts. Thus, there is evidence of a 
predominantly westerly wind regime in the Gawtet 
Ranges and surrounding area* during the period, or 
periods, of dune formation. 

There is, however, an anomaly between the prcsenr 
wind regime, as illustrated by the wind owe for 
Nonning (Fig 2b) and the presumed westerly wind 
of dune formation, since the winter sand-moving winds 
blow from the westerly sector, whereas the summer 
sand-moving Winds are from the SW, S and SE. It is 
presumed that most of the sand movement would lake 
place in summer endeT hot and dry conditions, with 
only minor transport in the moist, cool and vegetated 
winter conditions. But. if there were only a slight 
latitudinal migration of climatic zones during the period 
of formation of the dunes, as suggested tor example 
by MabbUtt (1977) and Spring (1979), then the region 



VanDel'R, W. t (LM85) Suhmcrgod dunes of northeastern 
Byre Peninsula MA Bicftfib University of Atfcluftlc 



(Unpub i 



would have heen influenced by summer rainfall 
maxima which would reduce sand movement during 
that season. On the other hand, dry winter conditions 
would be vuitabtc for the evidenced transport of sand 
by westerly winds. The lack of compatibility between 
dune orieruaiion and wind direction remains a problem. 
But assuming -a westerly wind regime, what factors 
are important in the formation of linear dunes 7 Why 
do parabolic dunes develop? How do the climbing and 
falling dunes form, and why do some of these form*, 
continue across the crests of the hills, whereas other; 
terminate on the upwind Ride only to resume on thr 
let- slope? J low aTc the transverse dunes formed? 

hntar duntr.s 

The origin of linear dunes j4 SfilJ debated (Cooke 
«f of. 1993 1. Where the dunes have been closely 
cwmmt'd, us in the Simpson Desert, these sand ridges 
appear to display the same range of morphology, and 
internal structures and temporal variations in 
asymmetry, indicative of formation under a 
bidirectional wind rep < me (McKee & Tibbetts 1%4; 
"Wopfner & Twidale 1967; Bnxikfietd 1970; Tseo 1990. 
1993} The linear dunes of the Oawlcr Ranges, 
however, developed in an upland setting rather than 
on desert plains. The confined valleys ought, in theory, 
to funnel the wind and hence to be conducive to a 
unidirectional wind regime, but bidirectional winds 
could be cither dominant or be superimposed on 
unidirectional effects. No structures have been observed 
within rhe dunes and, though this may reflect absence 
of deep exposures as much as any diagnostic factor, 
it * • not [visible to state whether the dunes have been 
shaped under a unidirectional or a bidirectional wind 
regime. 

Judging fn>rn die orientation of die linear dunes in 
tho Gawler Ranges, the airflow was apparently 
disturbed by the lulls of the province and was funnelled 
along valleys The hills induce zones of increased and 
of decreased air flow and of enhanced turbulence The 
dunes that are diverted around flanks of bills also reflect 
topographic control of the wind The changes in dune 
orientation and morphology between, on the one hand, 
the Great Victoria Desert, and, on the other, the 
Gawler Ranges, are due to several factors, First, tht 
westerly winds are diverted along the valleys. The 
linear dunes are not everywhere parallel with the 
regional air flow, as is eharaefcrastic of duneflelds on 
plains hut iheir orientation is. in part, determined by 
the local wind regime. Second, sand supply decreases 
within rhe upland where ihe silicic volcanic rocks 
weather less readily than do the granites to the west 
and. in particular, the supply of quartz grains is 
reduced. Third, sand movement is impeded as a result 
of the presence of near surface moisture, held either 
in valley alluvium or in rock fractures, and consequent 
vegetation growth 



DUVFS OF THE GAW1XR RANGES 



8 



Because of the lack of observed stroeuires in die 
dunes, the uncertainty about the relationship between 
dune morphology and wind regime arxl ihe tact that 
the dunes are relic and rww.sibiy related to different 
wind velocities, wind directions and rainlall amounts 
and distributions, the cJassificalBtJB d »hc dunes as 
linear, i.c eloujiate forms aligned in the direction <* 
the dominant sand moving winds, is tentative. 

Pcirabftta dimes 

The occurrence of parabolic rather than linear dimes, 
can he explained as follows In the Clorrohutnic 
Depression ro the S off the Gawler Ranges (Bourne ft 
dl. 1974) and elsewhere tMcKce l%6; Wasson tt of 
198^) parabolic dunes are located rn low lying aicas 
characterised by an abundani supply of sand and by 
proximity to groundwaters, which lead* to die luwei 
parts of the dune being stabilised by moisture *nd 
vegetation. This allows - the fngjier zones of sand to be 
transported downwind to give blowouts or 1 1 dunts. 
In the Gawler Ranges area the parabolic dunes occur 
only in wide open valleys and on piains. lot example 
in the western Moonaree Dunefteld and in prnvhe* in 
die Piccadilly DuneOetd. However, they arc not 
necessarily resu-icted to the towesl parts of thesc 
valleys. On the available evidence and as indicated nn 
the 1.100 000 topographic map with a contoui interval 
of 20 m. the WE belt of parabolic dutv-v 1»» »hc 
Moonaree Dunefield is sharply delimited on the 
northern side by a belt of linear dunes and , less sharply, 
on ihe southern side by dune-free plains. The pwrunoU 
duncs override low N-S uses in the valley and linear 
dunes occupy some low-lying areas in uV northern part 
of the durtefleld. Thus, in addition to stabilisation by 
vegetation and an abundant supply of sand, parabolic 
dune formation may require a critical wind velocity 
such as is attained only in wide valleys and on plain*. 

Climbing and fatting tluh*& 

The climbing ami (ahing dune* are a pomcubr 
variety ot linear dune which rise ;irul descend 
topographic obstructions wwnMtie li«eal wuhILs strong; 
enough to carry die available sand gram* np and over 
the topographic rises. The wind vrlnctiy is apparently 
reduced on approaching lite ptosfaclc arid deposition 
of sand occurs Many <tf uSe bounding slopes of the 
bornhardts are gentle (ar*YJL5-12 rt ) and reverse eddy 
How is generally not developed. And hence celw dunes 
(Tsoar 1983; see also Tig. 6a) arc not found windwanJ 
of cliffed obstacles. Where Ihe supply is dUffictettt, 
sand accumulaies until the dune reaches the height of 
the obstruction Where the bomuardt is low tin the 
Gawler Ranges <40 m) uV dune extends an to ami 
over the crest as a climbinje and falling duncv Where 
the bomhardt is high l>40 <*), the dune fotna may 
be discontinous though sand is carried on u> the ere*!, 
as evidenced by grains trapped in basins and cn?vi<rev. 



Downwind of the obstacle, however, there is a zone 
tif reduced wind velocity and sand deposition and dune 
formation occur- There may be fun her funnelling ol 
the sand to the lee of the obstacle where the falling 
dune is resumed in a topographic ernbayment (Fig. 6d). 
Ihe contrast between those linear tbrnis that continue 
Unbroken o v er crests of bedrock hills and those in 
which the climbing and Killing components arc 
separated, evidently reflects the Bernowlk effect (Pyc 
ft Boar 1990). 

Transit r\f dime:* 

The transveise dunes of the Scrubby Peak Dunefield 
occur immediately downwind of the Corrobinnte 
Depression, the presumed source of the sand, and 
where the sand supply is abundant. The bornhardts 
in this area stand about tCH.l m above the level of the 
plain, form N-S trending ndges and the bounding 
slopes are generally 5-10° with some as steep as 18°. 
I he plain is sand covered, with linear dunes of varied 
orientation, hut generally NW-SF where there are no 
topographic obstacles, N-S in the valleys and W-E on 
i he bornhardts rises (Fig. 4). Some of the N-S lineai 
dunes override topographic obstacles and hence are 
classed as climbing and tailing dunes. The W-F 
transport of sand across the bornhardt rises also 
explains the presence of sand In the valleys However, 
some of the dunes are limited to the upper slopes of 
fhe bornhardts and are located immediately to the lee 
of the crest of the bornhardts (Fig. 7). Although they 
may be linear dunes formed by winds from a northerly 
or southerly direction, in which case they do not 
conform to the pattern of dunes throughout the region, 
it is more likely thai these cresial dunes are transverse 
|m the originating wind it is suggested than the sand 
in these transverse dunes was driven up the windward 
slope of the bornhardts and little or no deposition 
occurred here due to acceleration of the air flow 
However, immediately downwind of the crest, 
separation of the air flow and deceleration occurred 
so that deposition of sand eventuated. However, further 
downwind, air flow accelerated and no dune formed- 

It is suggested thai in this area the dunes are a result 
of two different wind regimes, one a NW-SE wind mat 
was deflected by the topography and one a NW-SE 
wind dun was of sufficient strength to transport sand 
over the obstacles 

Signifa-ance vf lunettes and salinas in sand supply 

The lunettes located on the eastern shore of Lake 
Gairdner evidently spawn fields of linear dunes in their 
Ice in a manner similar to that described from the 
Simpson and other deserts (Twiddle 1972, 1981). The 
transport of sand to the Salinas by rivers and the 
formation of the lunettes arr important influences on 
sand supply and dune formation. Whether sand is 
curried by the wind from the W to the E shore of Lake 



\4 



D M, CAMPB£LL. C R TWIDDLE. S T ilUTTON * J K PRESC'CTT 



Gairdner (some 30 km) has not been determined. No 
dun$f have been observed on the bed of" the lake (J. 
Andrews, per*, comrn. 1994), though small barchanoid 
tonus have been reported on the bed of Lake Harris 
(R. Major, pels, codiid. 1992) Sand could be carried 
hy saJtation across the salina Given the hygroscopic 
character Of the halite crust this ma> be difficult to 
conceive, though Clarke (1994) described saJtation on 
some salrnas in Western Australia and S. Wells (pers. 
eotnni. 1994) has observed grains xalLttrng across a 
salt surface m California. Alternatively, sand reaching 
the W shore on the wind could be carried by wave 
action to the E shore during the occasional periods 
when there is water in die lake (Campbell 1968), 
though not from the lake bed unless the sail crust is 
dissolved or otherwise removed. Small ephemeral salt 
dunes, noted on the eastern shore of l^ake Gairdner. 
indicate the temporary redistribution of some tif the 
sail by deflation. 

Conclusion 



that the formation of parabolic as opposed to linear 
dunes is dependent on an abundant supply of sand and 
Fixing of the dune by vegetation only partially explains 
the distribution of these dune types in the Gawler 
Ranges province; olhei factors are apparently involved. 
Climbing dunes arc a variant of linear dunes and form 
in the zone of reduced wuid velocity upwind of an 
obstacle where the slope of the obstacle is gentle and 
does not generate reverse eddy flow. Falling dunes are 
associated with climbing dunes provided the sand 
supply is sufficient They develop in the zone ol 
reduced wind velocity to the lee of the obstacle. The 
crcstal transverse dunes are also due to deposition in 
the zone of reduced wind velocity Though the dunes 
of die Gawler Ranges area are essentially relic and are 
now stabilised by vegetation, there is sand movement 
during very high winds- The dunes were active about 
4000 years BP. The dunes of the Scrubby Peak 
Dunefield in the southern Gawler Ranges demonstrate 
that here the sand hits been transported by the wind 
at least 25 km from its source. 



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



Acknowledgments 

The authors thank J. A. Bourne for assistance in the 
field, V. A. Gostin for advice on sedinientology, K. 
Moxharn for advice concerning air flow around 
obstacles and R. Rice, R. Barrett and S Proferes for 
technical assistance. Two referees gave helpful 
suggestions on. an earlier draft ot the paper. 



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36 



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



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A REVISED SYSTEMATIC PLACEMENT FOR 

AUSTROTROMBELLA SOUTHCOTT 

(ACARINA: HYDRYPHANTIDAE) 



By MarkS. Harvey* 



Summary 

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

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



TfwwtitM* of thf ftnyal Snth-tx o/S. Aust. <t99f», 120(1). --7-4Q. 

A REVISED SYSTEMATIC PLACEMENT FOR AUSTROTROMBFLLA 
SOUTHCOTT (ACARJNA: HYDRYPHANTIDAE) 

by Mark S. Harvey* 

Summary 

Harvey. M. S. 1,1996) A revised systematic placement for AuxtroimmMlo Southcnii (Acarina: Hydryphantidae) 
front), K. Soc. 5. Aust 120(1). 37-40. 31 May. 199(5. 

Austrotmmbelfa Leprow Southeott, 1991. is transferred from the Trombeltidae iTrombidioidca) to the 
Hydryphantidae (Hydryphantoidea) and compared with other thymines of ihe Punisclhis gmup- 

Key Words; Taxonomy, Aeurina. Hydryphantidae .Austrotromhelto . Tromhcllidac. South Australia. 



Introduction 

The motiotypic genus AnsiwitornbeHa Southcott. 
1991, was recently described from four unusual 
specimens collected from wet shellgrit and soil beside 
the edge of a swamp near Robe, South Australia. The 
sole species, A. teuwsa. was extensively described and 
illustrated by Southcott (199J) and placed in the 
trombidioid family Trombellidae. However, 
examination of the type specimens lodged in the South 
Australian Museum (SAM), reveals that the gen.is is 
misplaced and more closely resembles water mites of 
the family Hydryphantidae than mites of the family 
Trombellidae. A redescription of the genus is presented 
here, along with an examination of its systematic 
position within the Hydryphantidae. 

Terminology mostly follows Cook (1974). 

Family Hydryphantidae Piersig, 1896 
Genus Aust rot rombella Southcott, 1991 

Aiwrotwmbelta Southcott, 1991: 207-208. 

Type species; /\uslrotrotntteila leprosa Southcott, 1991. 
by monotypy. 

Diagnosis 

Differs from all other mites by the following 
combination of characters: pedipalpal tibia with distal 
seta; swimming hairs absent: lateral eyes in capsules, 
idiosoma with numerous large plates: median eye 
present and situated near posterior margin of 
prefrontalia; three pairs of acetabula in anterior group 

Remarks 

Although regarded by Southcott (1991) as a merubcr 
of the trombidioid family Trombellidae, Attstrotrom- 
bflhi has more in common with the water mile family 
Hydryphantidae. In particular, the chelate morphology 
of the pedipalp, with a prominent dorso-distal tibial 
seta and a subdistally positioned tarsus, is virtually 

* Western Australian Museum Francis Street Perth v, Aust. 
6000. 



diagnostic tor the ramify (Cook 1974) and is completely 
unlike trombellids and other trombidioids which have 
the tarsus inserted subbasally on the tibia (e.g. 
Womcrslcy 1934). In addition, the idiosoma lacks the 
dense vestiture of setae characteristic of most adult and 
nvniphal trombidioids which is. instead, represented 
by longitudinal series of glandularia [termed cupolae' 
by Southcott (1991 )|. 

The presence of lateral eyes in capsules and the lack 
of swimming hairs places the genus within the 
Thvasinae (Cook 1974) and the large dorsalia and 
venlralia suggest a strong similarity with [he Pamseltus 
group as defined by Bader (1985). This group contains 
Panisellus K. Victs (with P. ihiermemanni (K- Victs) 
from northern Europe), Placothyas Lundblad (with P. 
uctoporu (K. Viets) from South Africa), Octothyas 
Lundblad (with O, hewintut Lundblad from South 
Africa), Poralhyas Lundblad (with P. thoracata Piersig 
and ft pritnittvu Lundblad from Europe and North 
Africa) and Thyasella K. V<ets (with 7. mandibuktris 
(Lundblad) from northern Europe}- Therefore, 
Austw/rombella and Us sole species, A. leprosa, is here 
transferred to the hydryphantid subfamily Thyasinae. 

Austrotrombella leprosa differs from these other 
genera in a number of small but significant ways. It 
closely resembles Panisel/us and Placotfiyas in the 
location of the postocularia within the prefrontalia and 
it differs from all members of the group by the 
possession of three pairs of acetabula in the anterior 
group (1-2 pairs in all others) and by the inclusion of 
the acetabula on to the genital flaps (published illustra- 
tions of all other genera appear to indicate that they 
are separate). It further differs from Panisellus by the 
presence of a median eye (absent in Panisellus) and 
from Plaavhyas by the posterior position of the median 
eye on the prefrontalia (situated near anterior margin 
in Placothyas) and the presence of 6*8 pairs ol 
acetabula in the posterior group (2 pairs in Pfacutliyas)- 

This species is only the second thyasine reported 
from Australia. The first, Notopanisus vinnulus Harvey 
from Tasmania, differs by the lack of large dor>alia 
and ventraiia (Harvey 19H8) 



ft 



M, S. HARVEY 



AustrotrombeUa leprosa Southcott, 1991 
(FIGS 1-8) 

.iustrotrombella leprosa Southcott, 1991; 208-211, Figs 
1. 2. 3a-e, 4a-c. 

Material Examined 

Holotxpe: 9, map reference (Penola 1: 250 000) 

283411, Robe district, S.Aust [37°12'S 139°47'E], 

in wet, alkaline, shellgrit-containing soil near swamp 

edge, under a stand of Leptospermum lanigerum 

(Aiton) Smith. 22.iii.1990, R. V. Southcott (SAM 

N1991112)- 

Paratvpes: 1 9, 1 o\ 1 deutonymph, same data as 
holotype (SAM N1991113-115). 

Diagnosis 
As for genus. 



Description of adult 

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





Figs 1-2. AustrotrombeUa leprosa Southcott, holotype 9- L Idiosoma, dorsal. 2. Idiosoma, ventral (setae omitted from 
one side). Abbreviations: dgl-6, dorsoglandularia; Igl -5, lateroglandularia; me, median eye; poo, postocularia; pro, prcocularia; 
vgl-5, ventroglandularia. Scale bar = 500 urn. 



REVISED PLACEMENT FOR AUSTRai'ROMBEUA 



39 



vg3 situated on level mid-way between genital flaps 
and anus; vg4 situated on same level as anus, vg5 
situated much closer to anus than to posterior margin 
of body (Fig. 2). Genital region (Fig. 3): genital flaps 
with setae on mesal edge and scattered over posterior 
third; 9- LI pairs of acelabula, 3 pairs situated in anterior 
third, remainder (varying from 6-8 per side) situated 
on posterior third, all acelabula circular. Chelicera 
(Fig. 7) of normal proportions, cheliceral claw curved, 
with several teeth; cheliceral lamella about two-thirds 
as long as claw, serrate. Capitulum without long, down- 
turned anterior extension. Pedipalp (Fig. 6): tibia with 
a thickened sub-medial seta on media] surface and with 
stoul distal seta. Pedal coxae covered with long, thick- 
setae (Fig. 2). Legs (Figs 4, 5) without swimming setae 
but most segments with numerous thick setae. Pedal 
claws completely smoodi (Figs 4. 5). Anus surrounded 
by ihick sclerotized ring (Fig. 2). 
Dimensions (/tm): holotype 9 ! body length 1464, 



width 1098; capitulum length 390; chelicera length 
367; genital field length 333, width 314; pedipalp; 
trochanter 63, femur 139. patella 112, tibia 195, tarsus 
51; leg I: trochanter 111, femur 250, patella 182, tibia 
236, metatarsus 269, tarsus length 255, width 64; leg 
IV; trochanter 250, femur 276, patella 179, tibia 378, 
metatarsus 380, tarsus length 300. width 52. 

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

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




Figs 3-8, Austrotrombetta leprosa Southeott, 3-7, holotype 9 3. Genital field- 4. Right leg L 5- Right leg IV 6. Right 
pedipalp. 7 Left chelicera. 8. Provisional genital field, paratYpe deutonymph. Scale bars = 200 /*m 3. 6, 7; 500 /xni 
4. 5; 100 um 8. 



AO 



M. S. HARVEY 



Description of deutonymph 

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

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



Acknowledgments 

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



References 



Bader, C. (1985) PomsMs-Studien: 6. Die Gattungen der 
Panisetlus-Gruppe (Acari, Actinedida, Hydrachnellae). Ent. 
Basil. 10, 7-17. 

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

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



Hygrobatidae and Athienemanniidae). Mem. Mus. Vict. 49, 

355-361. 
Southcott, R. V. (1991) A new trombellid mite (Acarina: 

Trombellidae) from South Australia. Trans. R. Sot: S. Aust, 

115, 207-212. 
Womersley, H. (1934) A revision of the trombiid and 

erthyraeid mites of Australia with descriptions of new 

genera and species. Rec. S. Aust. Mus. 5, 179-254. 



TRANSACTIONS OF THE 

ROYAL SOCIETY 
OF SOUTH AUSTRALIA 

INCORPORATED 



VOL. 120, PART 2 



SEVENTEEN NEW SPECIES OF CASTIARINA 
(COLEOPTERA: BUPRESTIDAE) 



By S. Barker* 



Summary 

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

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



Transaction* of the Royal Society of & Aust (19%). 120(2), 41-59. 

SEVENTEEN NEW SPECIES OF CASTIARINA (COLEOP1ERA: BUPRESTIDAE) 

by S. BARKKR* 

Summary 

Barker, S, (1996) Seventeen new species of Casmrina (Coleoptenr. Buprestiduc). Trvtts H. Sac. S. Ausi. 120(2) 
41-39, 31 May. 19%. 

Sevenieen new species ot Cuxtiarinu namely C. mlustti sp, nov.. C antarctica sp.. nov. , C aura sp. nov., C. 
(Kwrea sp nov., C charimk'ssa sp. nov., C, ititntn} sp. nov,, C tfamarzi sp. nov., C tmi^/mi sp. nov,, C fenupnta 
sp. BflV., C ht>tm;i>strm sp. nov., C jatkhnsenpHsrhi sp, nov,. C twnyma sp, nov., C nuthwboriva sp. nov . 
C* pauthnsatpuschi sp. nov.. C phucopus sp. nov., C suhewcta sp. nov., C ustuluut sp. nov., are described 
and three established species namely C t&flffl (Blackburn), C jetnumm (LaRme & Gory) C oiiospilota (LaRme 
& Gory) aiv redesenbed- 

Kfy Words: Colcopteru. Buprestidae. new species, Custiarina. 



Introduction 

The genus Castiurlna (LaPoric & Gor>) 
(Colcoptera: Buprestidae) is widely distributed in 
Australia and also occurs in New Guinea where ns 
distribution and abundance are virtually unknown The 
adults are often found on the (lowers of native 
Myrlaceae species and the larvae which are root and 
stem feeders are largely unknown, Although twenty 
two new Australian species have been identified 
recent!) (Barker 1993, 199?) a further close 
examination of material collected over many years has 
revealed even more new species. Ten of these ace 
associated with the Castiarina paraUcla (Wtyte) 
complex and all occur only in WA; two are associated 
with C uiiosptlota (La Porte &. dory) and one each 
of these occurs only in NSW and WA. The specific- 
status of a further species in this complex C femorata 
(LaPorte & Gory) is restored from synonymy: this 
species also occurs in NSW Five new species 
belonging to neither of these complexes arc described 
Ironi recently collected material 

The complex ut WA has previously been mis 
identified as C pamllda (White) but in (act this species 
occurs only in the eastern states. All memhers of the 
group are elongate and have dark pronouno and elytra! 
colours, most often rusty -brow n but dark blue in two 
species, a yellow margin and a .-.ingle row of yellow 
spots along the middle of each elytron with a maximum 
ot lour in each row and minimally one. when all of 
the spots are fused. All species in the complex have 
a dense layer of silver, flattened, feathery hair lining 
the entire ventral surface and encroaching on to the 
lateral surfaces of the pronotunt- This dislingu t 
them from all other Custiarina which have thin. roanO 
ban on '.he vcnlrul surface Most o\' the new specie 



Department of /oology. University ot Adelaide S 

safe 



also have sculptured proetigers in both sexes and this 
feature is unique among Custiarina as all species 
outside this complex have rounded, unsculptured 
proeUgers. The two species previously described from 
the complex in WA are C cnnvcolor fl .a Forte & Gory) 
and C oaoptwcnifa (Barker 1995). Because this is the 
most difficult group of Castiarina to identify, 1 have 
included a key to the twelve known WA species. 

C. ocrospihra (LaPorte Ac Gory) has a dark head 
with a yellow frontal spot, dark pronotum with yellow 
lateral margins and dark elytra! markings with yellow 
spots. The ventral surface is yellow with blue sututal 
markings and blue legs. A species occurring on the 
eastern edge ot the Darling Scarp, WA and on the 
coastal plains has long been misidentificd as C, 
oaaspilnia [t resembles C rinrra (Blackburn) which 
occurs inland but is easily distinguished by differences 
in colour, being blue and yellow with blue legs whilst 
C rinrta has some red markings on the elytra and red 
legs With red sulura! markings. The aedeagi are 
tiiffcrenl . There appear to be two species, C. cituta 
which t.s redeseribed and a new species which is 
described. 

None of the remaining five species ts close lo each 
other and all arc distinctive. They have all been 
collected recently, one of them by use of a lure, a 
technique not used before for the capture of Castiarina. 



Materials and Methods 

Male genitalia were prepared and illustrated by the 
method described by Barker (1987). The holotypc is 
illustrated in all new species except one in which the 
allotype is illustrated Measurements given are mean 
total body length and width with standard error, except 
where there arc insufficient specimens to make the last 
.Ail-jl.tOon. Codens used in the trxt far museum and 
private collection* following the four letter .system of 



<\2 



S. BARKER 



Watt (1979) and Araett et at. (1993) are: ANIC 
Australian National Insect Collection, Canberra 
BMNH; British Museum (Natural History). London 
MNHN; Museum Nalionale Histoire Naturelle, Paris 
NMVA: National Museum of Victoria, Melbourne 
SAMA; South Australian Museum, Adelaide; WAMA 
Western Australian Museum. Perth; HDWA Mr H 
Demar?. Guilderton; JHQA: Mr J Hasenpusch 
Innisfail; MHSA. Mr T. M. S. Hanlon. Sydney 
MPWA: Mr M. Powell, Melville. 



Castiarina jemorata (LaPorte & Gory) 1838 
(FIGS IB. 2B) 

Stigmodera femoraut LaPorte & Gory 1838: 37, PI 
8, Fig. 42. 

Stigmodera (Casriarina) oetnspilota var. roseipc* 

Deuquet, 1956 (new syn.). 

Hole/type: Sex unknown, S. femnrata LaPorte & Gory, 
Aust. MNHN (not seen) 




Fig. I. Photomicrographs of aedeagi and male and female proctigers of the following Casiiarittti specie*. A. Custiahna 
damn) sp. nov. B. C. femorata (L & G). C. C. cincta (Blackburn). D. C subemcta sp. nov. E. C octaspiloia (l. & G). 
F C. paulhiisenpitschi sp. nov G. C. bucolica (Kerremans). H. C. demarrjsp. nov. I. C. placida (Thomson). J. C. nullarborkxi 
sp. nov K. C tydista (Rainbow). L. C jackhasenpuschi sp. nov. M C ablaut Barker. N. C. hemizostera sp. nov. O. 
C antarctiat sp. nov. I. proctiger male. 2. proctiger female. P. C nrtoptwrtata Barker. I. proctiger male. 2. proctiger 
female. Q. C. ferruginea sp. nov. 1. proctiger male. 2. proctiger female. R. C, adusta sp. nov. I. pnicliger male. 2. proctiger 
female. S. C. aura sp. nov. 1. proctiger male 2. proctiger female. T. C chariemessa sp. nov. I. proctiger male. 2. proctigci 
female. U. C. uzurea sp. nov. 1. proctiger male. 2. proctiger female. V. C ustutata sp. nov. 1. proctiger male 2. proctiger 
female. W. C ptuieopus sp. nov. 1. proctiger male. 2. proctiger female. X. C. nonyma sp. nov. I. proctiger male. 2. proctiger 
female. Y. C enigma sp. nov. f proctiger male. 2. proctiger female. Z. C. erucieuhr (L & G). I. proctiger male. 2. 
proctiger female. 



SEVENTEEN N£W SPECIES OF CAS7URINA 



43 



Colour 

Head dark blue with green and purple reflections, 
elongate yellow frontal spot, muzzle blue, Antennae 
blue. Pronotutn dark blue with yellow lateral margins 
w idth increasing basally. Scutellum dark blue. Elytra 
yellow with following dark blue markings: narrow 
basal margin, elongate sinuous vitta on each side 
meeting basal margin over humeral callus, meeting 
post-medial lascia at margin enclosing spot on margin. 
apical mark enclosing elongate yellow mark between 
it and posi-medtal fascia, small yellow apical spot on 
each side of suture, marks all connected along suture. 
Ventral surface yellow with testaceous-red sutures arid 
testaceous-red edges to abdominal segments, except 
S-. Legs; coxae and trochanters testaceous- red and 
dark blue: femora mainly testaceous-red, apiwlly dark 
blue, libiae a dark blue with ventral testaceous-red 
mark near ventral apex: uirst blue. Hairs silver 

Shafh' and sculpture 

Head shailowly punctured, median sulcus natrow. 
nwz?Je short. Antcnnomeres 1-3 obcomc, 4 hall 
toothed, 5-11 toothed, Pronotutn shailowly punctured, 
apical margin straight, basal margin barely bisinuate, 




Fig.2. Habitus illustrations of the fallowing Castiarvui species. 
A. Castiarina nctospiloUi (L& G), B C femurum (L & 
G). C. C. Mibcincta sp. nov. holotype. D. C danmj \p, 
nov. holotype E. C cinaa (Blackburn). 



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

Sitf 

Males. 12 4 ± 0.35 x 4. u ± 0.14 mm (20). Females, 
114 ± 0.22 x 55 ± 0.09 mm (5). 

Aedeagus (Fig IB) 

Parameres angled outwards from basal piece, 
tounded premcdially then narrowed, rounded apically 
Penis sharp, sides acutely angled away. Hypophysis ol 
basal piece medium width apices rounded. 

Distribution 

NSW: Armidale district, central coastal . 

Remarks 

This species was synonymtsed with C. QClOtipUoW 
by Saunders (1868) who was followed by all subsequent 
authors Deuquet (1956) gave the varietal name rosnpes 
10 a specimen he identified as S. octospitota. A single 
male specimen in the South Australian Museum idcn 
olied as Srig.S-spilota L & G var. roseipes Ucuq. in 
Deuque.t's handwriting, is clearly a separate specie? 
front C octospitota. The holotypes of C octospitota 
and C.fcwomta are both odged in the MNHN but 
cannot be identified because their labels, along widi 
those of all other LaE*orte & Gory type labels, havtr 
been removed. Dcuquet's description of the red femora 
and fed vermiculation on the ventral surface of his 5, 
octospitota var. roseipes concurs with the original 
description of S. femorula. The figure of $ femorata 
/LaPone & Gory 1838, PI 8, Fig. 42) also conforms 
in general, except that the illustration shows the base 
of the head to be yellow instead of the yellow frontaJ 
spot in the Dcuquet specimen. J assume that this i-. 
artistic licence on the part of the illustrator because 
none of the species in this species group has a head 
with a yellow base. In the figure the pre-medial fascia 
is complete. A similar pattern is present in only two 
of the fourteen specimens examined. All other 
specimens have the pre-medial fascia incomplete; thus 
two spots on each side of the suture coalesce to form 
a sinuous yellow vitta. S. oaospilota var. roseipes 
Deuquet is undoubtedly a svnonvm of C femorata (L 
&G). 



n 



vS- BARKER 



Casuannu octospilota (LaPOrte & Gory) 
(FIGS IE. 2A) 

Stigmodem oaospilota UHorte & Gory 1838: 28, Fig. 
29, Pi. 6. 



Distribution 

NSW: Blue Mrs, Sydney, central 10 north coastal. 
Old- Southern and central coastal, Blackdown 
Tableland, Shrove Is. 



ifijhtypc Sex unknown, Australic, MNHN (not seen) 

Colour 

Head; muz/Je blue bnse dull green- purple; yellow 
frontal mark. Antennae blue. Pronolum: medially dull 
green-purple; laterally yellow, base wider than apex, 
Scuteilum black with dark blue reflections. Elytra 
yellow wiih following dark blue markings: narrow 
basal margin, small and narrow pre-medial fascia 
absent in many specimens, connected to long oblique 
vitta from lower end reaching lateral margin, enclosing 
very large basal yellow spot and smaller one on margin, 
btoad post -medial fascia reaching lateral margin, 
slightly angled posteriorly, enclosing large yellow mark 
between u and first fascia; mark covering whole ape*. 
enclosing small yellow mark between it and second 
fascia Ventral surface yellow with blue sutures. Legs 
blue, flairs silver 

Shape and sculpture 

Head closely punctured, median sulcus small, 
muzzle short Antennomeres 1-3 obconic 4 half- 
toothed, 5-11 unnhed Pronolum closely punctured, 
minute basal fovea extending forwards to middle as 
glabrous line, basal notches represented by glabrous 
areas, more marginal than medial; apical margin 
straight, basal margin bLsinuate; laterally rounded out 
from base, widest pre-medially, rounded and narrowed 
to apex. Scuteilum scuofarm, glabrous, excavate at 
basal edge. Elytra puiietate-striate. intervals convex and 
punctured, less so medially than elsewhere, laterally 
angled out from base, rounded at humeral callus, 
concave, rounded post-medially and narrowed to 
tmpinosc apex; small marginal and medial spines, 
margin between straight, widely separated, small 
sutural spine, close to medial spine, apices divergent 
Ventral surface with shallow punctures, edges of 
abdominal segments glabrous, elsewhere moderately 
hairy, hairs medium length S 7 truncate and indented 
medially in both sexes 

Size 

Males. 12.5 ± 0,28 *5J ± 0.20 mm (33). Females, 
U6 ± 0.23 x 56 ± 0.11 mm (40). 

/tedeagus (Fig. IE) 

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



Castiannu <tncta (Blackburn) 1890 
(FIGS IC. 2Jz) 

Stiptnodera < incut Blackburn 1890: 13, 157 
{replacement name tor S. rubrocincta Kerrenvans 
1890: 46 primary homonym 5. ruhrocmcta Gehin 
l:s55) 

Holotypc- 9, £ fubfoaucra Kei remans. Boucard 
Australie_ BMNH 

Colour 

Head: muzzle blue-green; base purple-green; large 
yellow frontal mark. Antennae blue-green. Pronolum 
laterally yellow, medially purple-green. Scuteilum blue 
Flytra yellow with the following dark blue markings 
basal margin; prc-medial fascia not reaching margin 
with ends projecting anteriorly to basal margin as vitta 
enclosing large yellow basal spot, elongate yellow mark 
on margin ;it humeral callus, narrow red apical margin 
p0&] -medial lascia not reaching margin, enclosing 
yellow band between it and first fascia, prc-apical mark 
HI shape of short fascia enclosing elongate yellow band 
between it and second fascia, all marks connected along 
suture. Lateral red margin on the two intervals from 
humeral callus, broader at pre-apex and apex Ventral 
surface yellow; sternum lateral blue-green sutural 
marks, red medially and along edges v>f abdominal 
segments. Legs: femora blue-green apically, red 
medially, tibiae and tarsomeres blue-green Hairs 
silver, 

Shape utut sculpture 

Head with shallow punctures, flat, muzzle short 
Antennorneres 1-3 obconic, 4 halt-toothed. 5-11 
t>»olhcd. Pronolum with shallow punctures, basal fovea 
represented by shallow depression, very small basal 
notches more marginal than medial; apical margin 
straight, basal margin almost straight. Scuteilum 
setuifbrm, without punctures, e<cavute along basal 
edge. Elytra punctate-stnate, intervals flat medially, 
convex apically and laterally, punctured, less so 
medially, laterally angled out slightly from base, 
rounded al humeral callus, concave, rounded post 
medially and narrowed to tmpinosc apex; marginal 
spine small and sharp, median spine larger and blunt, 
sutural spine small and sharp, margin rounded between 
spines. Ventral surface with shallow punctures sparse 
short hair. S ? : truncate both sexe*. 



SEVENTEEN NKW SPECIES OF CASTIARtNA 

Mates, 11.3 dt 0.22x 4.3 ± 0.10 mm (24). Females 
12,2 ± 0.34 \ 4.7 ± 0.15 mm (20). 



45 



on each side. Ventral surface orange-yellow with blur 
marks along sutures and along edges of abdominal 
segments. Legs: femora and tibiae blue; tarsi brighi 
blue. Hairs silver. 



Aedeagus (Fig. IC) 

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

Distribution 
Occurs in inland south- western WA. 

Casfiurina darattj sp. nov. 
(FIGS IA, 2D; 

Hohtvpe: a. 4km NE RocWy Glen, NSW, 3.xi.l98I, 
S. Barker. SAMA J 21 300. 

Allotype; 9 . same data as hololype, SAMA I 21 301. 

ramtvpes: NSW; 1 9. 43 km S Narrabn, 27.x 1975. 
S. Barker, SAMA; 14 c o\ 8Q 9, Binnaway, 
2.\i 1981, S. Barker, SAMA. I 9. 6 km SW Rocky 
Glen, 3. xi. 1981. & Barker, SAMA; 2 era, 2 9 v- 4 
km SW Rockv Glen, 3 vi.1981. S. Barker, SAMA. 1 9, 
3 km SW Rocky Glen. 3xi.l981, S. Barker. SAMA; 
|o\ 1 9,2 km SW Rocky Glen. 3 vi .1981. S Barker, 
SAMA; 5ry o\ 49 9 ( same data as holotype, SAMA, 
2cr '.y , 2 9 9 . Garrawilla TO 6 km NE Rocky Glen. 
3.X1.198I, S. Barker. SAMA. 2 9 9. Garrawilla TO , 
8. u. 1981. R. Anderson, SAMA; I 9. Garrawilla TO., 
12. xi. 1981. S. Barker. SAMA; to*. 6 km Nfc 
^uabarrahran, 6.XU983, A. M. Sundholm. MHSA; 
30" ct % 1 9. 40 km 6 Coonaharrabran, 9 xi.1990. T. 
M. £ Hanlon, MHSA. 1 o\ 60 km N Coonabarmbran, 
9 xi.1990, T. H. S. Hanlon, MHSA, 2a a, 29 9. 
40 km E Coonaharrabran. 8 xi 1991. T M. S. Hanlon. 
MHSA; 1 cr, Rimnd Hill. 21 xi.l99L T. M S Hanlon, 
MHSA; Ur, 30 km H Parkes, 29 xi. 1993. T M S. 
Hanlon. MHSA. 

Colour 

Head black with given reflections, mu77.ie blue, large 
orange-yellow frontal spot. Antennae blue. Pronotum 
medially dark blue, laterally orange-yellow increasing 
in width basally. Scutcllum dark blue. Elytra orange- 
yellow with the following black markings with blue 
reflections; narrow basal margin: sinuous vitta from 
basal murgin over humeral callus meeting margin and 
suturai mark enclosing yellow -orange elongate mark 
on margin and basal spot; bmad post-medial fascia 
reaching margin enclosing large orange-yellow mark 
between it and first mark; mark covering apex mating 
post-medial fascia on margin and enclosing a large 
orange-yellow mark; small orange-yellow apical spot 



Shape and sculpture 

Head shallowly punctured, median sulcus shallow, 
muzzle short Antennomercs 1-3 obeotiic. 4 half- 
toothed. 5-11 toothed- Pronotum shallowly punctured, 
narrow basal fovea extending forwards to middle as 
glabrous line; apical margin projecting medially, basal 
margin barely bisinuatc; laterally parallel-sided al base. 
rounded to apex. Scutellum scuufonn. glabrous, 
excavate. F.iytra punetate-stria'e, intervals convex, 
lightly wrinkled and punctured; laterally angled out 
from base, rounded at humeral callus, concave, 
rounded post-medially and narrowed to trispinose 
apex; spines small, margin straight between marginal 
and median spine, rounded between median and suturai 
spines, apices divergent. Ventral surface with shallow 
punctures, edges o\ abdominal segments glabrous. 
cUewherc moderately haiiy. sparse medium length 
hair S r male truncate, slightly indented medially; 
female truncate, deeply indented medially, margin 
overhanging apex which is covered with bristles. 

Sfee 

Males. 10.4 * 0.15 x 4.0 * 0.06 mm f34). Females. 
II 1 * 0.20 x 4.4 i 0.09 mm (26), 

Aedeaxus (Fig. IA) 

Short. Parameres angled outwards from basal piece, 
rounded pre-medially, parallel sided, apieally rounded. 
Perns sharp, sides acutely angled away Hypophysis Cjf 
basal piece medium width, apical! y rounded. 

Remarks 

The basal colour of this species fades rapidly in death 
from orange-yellow to pale yellow. Both C octospitotu 
and C. femoruia have a yellow basal colour in life. Also 
it is smaller than the other two species and the male 
genitalia are smaller and a different shape (Figs IA. 
IB, IE), 

Etymology 
Name derived from Arabic damn], orange. 

Qxstiarina subcincfa sp. nov. 
(FjgS ID, 2Ci 

Holotype: if t Bold Park, City Beach, WA, 3 xi.1976, 
R P. McMillan. SAMA I 21 302. 

Allorvpe; 9, City Beach, WA. 3.xii 1955, J. A. L. 
Watson, SAMA I 21 303. 



46 



S. DARKER 



Pararypes; WA. L9. Cannington, 12 xii.1954, S, 
Barker, SAMA; 3o-o% City Beach, 24.x, 1954. S. 
Barker. SAMA; 12o*o\ 3 9 9. City Beach 
26.xi.25.xii. 1955, J. A. L. Watson* SAMA; 1 9, 9.5 
km SW Jarrahdale, 11 \i 1956. S. Barker. SAMA; 1 o\ 
City Beach. 6. u. 1957, S. Barker, SAMA; la, City 
Beach, 2 x 1964, S Barker. SAMA; ?o-o\ Wembly, 
3.x. 1970, S. Barker. SAMA, 4cro\ I 9. same data 
as holorype. SAMA. 2o , 0\ J 9, Glen Eagles, 
7.1.196% S Barker* K. T Richards. SAMA. Zcrcr. 
3 9 V^ Walyunga N P., 4.xi.l984, T. M S Hanlon, 
MHSA; 5cr cr, 1 v, Wembly. 4.xi.l9S5, T. M. S. 
Hanlon, MHSA; J o\ Swanbourne. 23. a 1991. T. M, 
S. Hanlon, MHSA, 6c'C, 19, Swan R. H. W. 
Brown, SAMA 

Cutour 

Head nasally dark blue with green and purple 
reflections, muzzle blue, large yellow frontal spot 
Antennae dark blue Pnvnotum medially dark blue with 
green and purple reflections, laterally yellow. Scutellum 
dark blue. Elytra yellow with following black markings 
with blue reflections; narrow basal margin; sinuous 
vitta from basal margin over humeral callus meeting 
narrow pre-medial fascia close to margin enclosing a 
yellow spot on margin and large yellow basal spot; 
bfuad post-medial fascia reaching margin enclosing 
yellow spot between it and pre-medial fascia, mark 
covering apex enclosing elongate yellow mark between 
Jt and post-medial fascia and variable ,*pital yellow 
spot, all marks connected along suture and along 
margin except at humeral callus; outer margin of apical 
spot variably red, Ventral surface yellow, sutures blue 
and lateral blue spots on S 4 . S v S f . Legs blue. Hairs 
silver. 



Size 

Males, 12.8 ± 0.13 x 4.9 ± 0.06 mm (41). Females. 
13.5 ± 0.22 x 5.2 ± 0.09 mm (24). 

Aedeagus (Fig ID) 

Parameres angled outwards from basal piece. 
rounded pre-medially. parallel-sided, rounded apically. 
f^nis sharp, sides acutely angled away. Hypophysis rif 
basal piece narrow, apically rounded 

Xewarky 

This species, previously confused with t 
ortospilota, forms a species pair with C anctu 
(Blackburn). It occurs on the coastal plain of WA and 
on the western edge of the Darling Scarp whereas C 
cmcta occurs in the more arid inland qt$AS oi tin south- 
west. It differs from that species having only very small 
red markings on the elytra and not »m the leg-, ui 
abdominal segments C, cineta has red markings on 
the elytra, red femora and red sutures on the ventral 
surface. Also the elytra! spines are more obvious in 
C. suhcincta than in C 1 inrta and ibe male genitalia 
are a different shape (Figs 1C. ID). 

Etymology 

The name i&denvcd (torn L nth, under. I. rinrium, 

girdle. 



Ctistuuina adu&ta sp. nov. 
fFigs 1R, 1 Rl. I R2, 3J) 

Holonpc o\ 5 km W Mt Dale. WA. 13 s 1V80. S. 
Barker, SAMA 1 21 304. 



Shape and sculpture 

Head shallow!) punctured, median sulcus small and 
shallow, muzzle short Antennomeres 1-3 obeonic, 4 
half -toothed, 5-11 toothed. Pronotnm shal lowly 
punctured, narrow basal fovea extending forwards n» 
middle as glabrous line, basal notches represented by 
glabrous area on each side closer lo margin than 
middle, apical margin projecting medially, basal 
margin almost straight, laterally parallel-sided at base. 
rounded from base to apex. Scutellum seutiform, 
glabrous, flat. Elytra punctate-.striate, intervals convex, 
wrinkled; laterally angled out from base, rounded at 
humeral callus, concave, rounded post-medially, 
narrowed to trispinose apex; small marginal spine, 
larger medial spine, smaller sutural spine, margin 
rounded between spines, apices diverging Ventral 
surface with shallow punctures, edges of abdominal 
segments glabrous, elsewhere sparse medium length 
hairs. S 7 truncate both sexes. 



&tkm& 9 ■ Uke Grace, WA. I9.X.1970, K & E, 
Carnaby, ANIC 

Pnraivftes: WA 5 n- o*. i y < T \ same data as allotype . 
ANIC, 1 V , 80 kin E Hyden. 29 x 1984. M Powell, 
MPWA. lo-; 1 km WNW Bonnie Rock, 20. k 1990, 
S. Barker, SAMA 

Head, antennae and pronotum bronze, Scutellum 
dark blue. Elytra yellow with the Ibllowing brown 
markings: marks coalesced leaving a continuous yellow 
margin from base to near apex , a row of four elongate 
ftpdtS down each elytron, the first two variably 
connected- Ventral surface and legs bronze. Hairs 
silver. 

Shape and sculpture 

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



SEVENTEEN NEW SPECIES OF CASTIARINA 



47 




Fig. 3. Habitus illustrations of the following Casiiarina 
species. A. Castiarina nonyma sp. nov. holotype B. C 
rrartcotor (L Si G). C. C. enigma sp. nov. hololype^ D. 
C, tmtarctica sp. nov. holotype. E. C. phaeopus sp. nov, 
holotype. F. C. ustidata sp. nov. holotype. G. C. azurea 
sp. nov. holotype. H. C. ferrugtnea sp. nov. holotype. ). 
C aura sp. nov. holotype. J. C. adusta sp. nov holotype, 
K. C charientessa sp. nov. holotype. 



meres 1-3 obconic. 4-11 toothed. Pronotum shaJlowly 
punctured medially, larger and deeper punctures 
laterally, narrow basal fovea extending anteriorly to 
middle as glabrous line, basal notches represented by 
glabrous area on each side closer to margin than 
middle; apical margin projecting medially, basal 
margin barely bisinuate; laterally parallel-sided at base, 
rounded and na r rowed to apex . 1 ateral ly hai ry. 
Scutellumscutiform, punctured, flat. Elytra punctate- 
striate, intervals convex, wrinkled and punctured, 
laterally angled out from base, rounded at humeral 
callus, concave, rounded post-medially and narrowed 
to bispinose apex; spines small and blunt, margin 
variably rounded and indented or straight between 
spines, apices hardly diverging. Ventral surface with 
shallow punctures, edges of abdominal segments 
glabrous, elsewhere hairy, hairs llatlened and feathery 
Legs: femora hairy with flattened hair. S 7 ; males 
truncate; females rounded. 

Ske 

Males, 14.5 ± 0.12 x 5.0 i 0,06 mm (7), Females. 
15-9 =b 0.45 x 5.4 ± 0.12 mm (5) 

Aedeagus (Fig. 1R) 

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

Female tenmntdia (Fig. 1 R2) 

Proctiger with apical edge flattened, rounded 
laterally. 

Remarks 

C. adusta sp. nov. is the largest member of this group 
in WA. It can be separated from C. jerrugineu sp. nov . 
the next largest brownish species, by its si?c, the 
conformation of the elytral markings - there, are lour 
spots in C. fenugineu and three in C. adusta, its 
relatively unsculptured proctiger in both male and 
Icrnalc. whereas both sexes of C femiginea have 
bilobed proctigers and in females they arc spined The 
aedeagus in C. fitrrugihea is broader at die apex than 
that of C adusta (Figs IQ> 1R) 

Etymology 

The name is derived from L adustus, brown. 



Castiarina azurea sp. nov 
(FIGS 1U, 1 UI, I U2, 3G) 

Holotype: cr , 2 km £ Tallering Station, Pindar, WA. 
22.ix J989, S. Barker, SAMA 1 21 305. 



i!-; 



S. BARKER 



.\il<>tvp<-: v , same data as holorype, SAMA r 21 30ft. 

Puratxpes- WA: 1 cr. Goomalling, 13. ix .1953, R. P 
McMillan. WAMA; L cr, 2 9 9> Moorine Rock. 
16 x ISS3, F. H. Uther Baker, SAMA, 19. Wialki. 
9. 1 a. 1957, S. Barker, SAMA; 1 cr, Toolibin, 18.x 1958. 
F H Uther Baker, SAMA; Jlcyp*, Burraeoppiu. 
16x4963, F H. Uther Baker. WAMA; 3 cro\ 2 9 9- 
78 kmNE Wubin, I7.ix 1970. S. Barker, SAMA; I 9, 
9$ km NE Wubin, |7.ix|970, S, Barker. SAMA; 
l&®, 1 9. 55 km S Paynes Find, 18. iv 1970 SAMA; 
l ti I v - s 7 km S Paynes Find. lH.rx.1970. S. Barker. 
SAMA; 2 9 9- W km E Elacbbuning Rock. Wialki. 
211 ix 1970. S. Barker. SAMA; 5a- o\ 29. Wialki. 
31. ix 1970, S Barker, SAMA: 2cro\ 29 9. 
WulgooUn, 9ix.l97l. F H. Uther Baker. SAMA, I 9. 
Tallenng Station* Pindar. 3,ix 1976. R. P. McMillan. 
SAMA. 5r/o\ 3 9 9, BMl kW WSSV Cnolgardie. 
lKix 1976, R J Chrnnock. SAMA; I cr, 50 km N 
K.ilh;..rn 20vui.t9?8 ( M fowctl, WAMA; I 0\ Ballinc 
.Siai.-.n, 24/25.VH.7979, A. M. & M. J. Douglas 
WAMA; lor. 2 9 9 . Muckinbudin, 10.U979. R. P 
McMillan, WAMA; cr, 16 km E Mt Magnet 
20 .1 K 1980. S. Barker & D. I Williams. SAMA: I ct 
19 km N Carnarvon, 22. ix 1980, S. Barker & a J 
William*, SAMA; Iff, 2 V 9, 89 km N Carnarvon. 
22 ix 19X0, S Barker & D. J. Williams, SAMA: 1 cr 
] ? . 44 km B Kalbarri. 26. i\ 1980. S. Barker & D. 
J. WlBlattM, SAMA, 3crcr. 46 km B Kalbarri, 
l$fo 1980. 5 Barker <fc D, J. William. SAMa.2o-o\ 
19. S Ram paddock, Tillering Station. Pindar 
27 .ix. WW S, Ba/kerA D. I Williams, SAMA; let p , 
i v 17 km W Mullewa. 29 ix. 1980, S Barker & D. 
J. Williams SAMA. 29 9- Mt Walker, 23 .x.l98tt R 
P Mt-Miil-.m. WAMA; I y, Cubhin, 29.x. 1981. R, P 
McMillan. WAMA; 2 9 9- Southern Crass, x 1981, 
R P. McMillan, WAMA: IjOp, 39 9. 2 km N 
EvflQ5ton< 23 h 1982. B. Haitich & I. R Houston, 
WAMA, J 9, 64 km NE Esperance, 18x1982. S 
Barker, SAMA, ler. Bullfinch, 2.x 1983, B Jones, 
MPWA; 19 35 km W SaJmon Gums, 8.x .1983, G. 
Browning & G. Mutze, SA M A . 3 cr o\ ) 9 , Southern 
Cross 8Xl9(ft K P. McMillan. WAMA. Geo*. 
'J v 5 1 Esperance to Norseman Hwy. 35 km W T.O 
to Pcflke Charles 9. K 1983. G. Browning $ G. Muue. 
SAMA. 29 9. 35 km E Merredin. 24.x. 1983, G 
Browning, SAMA; 1 a\ 2 9 9 , 50 km E Merredin, 
24. \. 1983. SAMA. I CK, Ubethn Rock, I6.ix.l984. R 
P McMillan, WAMA; Icr, J36kmNE Payne's Find. 
30. ix 1984. M. PmveH. MPWA, I -cr, 3 kill W 
Dowcnn. 22\ 1984. R P McMillan. WAMA; ICf, 
Eneabna, 4 x.1985. R. P. McMillan, WAMA; 1 o\ N 
Tann Rock reserve, 15/16. x.1985 T. F. Houston. 
WAMA. IO 75 km B Hyden. 24/27.X.I985. T R 
Houstoti. WAMA; 6o» cr, Woi Coorow, 2.x 1986, A. 
G. WetK WAMA; 1 cr, 16 km NE Mcrredm, 9.x >986, 
R P McMillan. WAMA; fcTOr. Encabba. x.1986. R. 



P. McMillan, WAMA; Icr. Bindoo Hill reserve. 
I2.ix.1987. T. F. Houston, WAMA; 2a cr. 1 9, 7 km 
SSW Jiugemarra Station. 24/26,viii.l988. R. H 
McMillan & T F. Houston. WAMA; 1 9 . Shark Bay, 
29.viii.I988, A. Hay, MHSA: Icr, 110 km N 
Carnarvon 18 ix 1989. S. Barker. SAMA; ley o\ I 9 , 
Pindar paddock. Tallering Station, Pindar, 2l.lx.i9S9, 
S Barker, SAMA; 7&CT, same data as holotype, 
SAMA; I cr , I 9, 3 km N Tailoring Station. Pindai. 
22.tx.l989, S. Barker, SAMA; 2a cr. 6 km N Taflerinu 
SUIion. Pindar. 22.ix 1989. S. Barker. SAMA Si -' f 
V) km N Tailoring Station. Pindar, 22. is 1989, S 
Barker, SAM A ; 1 o* , gravel ha\ t Bonnie Rock. 
20. ix 1990, S. BarkCti SAMA. 1-CPC^ Merredin. 
21 K.BB1. 7. M, S. Hanlon. MHSA; SCO 1 , Ghooli, 
21 ,1991. T M S. Hanlon. MHSA. 2crcr. 3 km S 
\eilwline.2Lx.l991. T. MS. Hanlon, MHSA 5a- cr, 
N7T, 3: km E Southern Cross, 21x1991. T. M. S 
»-'onlon f MHSA. Icr. Queen Victoria Springs. 
2i::.Ml992. O. Knowlcs, MHSA; 2<xrr . IQ, 
Noriham, SAMA; Jxftfi AnkerteU. H. W. BroWD 
SAMA. 

CeJtHtr 

Head dark blue with purple reflections Antenna* 
bronze. Pronotum dark blue wtLh purple rcflecrinns 
Sctitellum purple. Elytra with yellow background 
colour and dark blue elytra! markings, coalesced 
forming four yellow spots nn each elytmn. basal (rtora 
or less rounded, pre-medial elongate, post metlial 
rounded, pre apical elongate, basal and pre-medial 
coulcsced in about half specimens examined lornting 
aJi elonyate mark, ptist-medial and pre-apical coalesceil 
in only one ypecinien examined, yellow margin from 
base to near apex VentTgl surface bronze wiu/i coppfitj- 
parple reflecrionx Haos Mhet 

Skami and siulpinre 

Head closely punctured, median sulcus present, 
Diu^e .short. Arucnnomcres 1-3 obcome, 441 (oothe^ 
Pronotum closely punctured, narrow basal fove;i 
extending forwards to middle as glabrous line, basal 
notches represented by glabrous area on each siJr 
closer to margin than middle; apicaJ margin projecting 
medially, basal margin almost straight; laterally angled 
inwards from base for short distance then angled 
outwards and rounded to widest part post-medially 
loundod and narrowed 10 apex Seutellum seuurbrm. 
punctured, flat Elytra punctate-sinute, intervals 
convex, wrinkled and puncma-d. more heavily lateralb 
than medially; laterally angled out from base, rounded 
at humeral callus, concave, rounded post medially, 
narrowed to btspinose apex; marginal spine small and 
sharp, sutural spine minure, margin indented jnd 
rounded between spines, apices hardly diverging. 
Ventral surface with shallow punctures, edges ul 
abdominal segments glahrous. elsewhere hairy, hatrs 
medium length, flaltened and lealhery. fL: truncate in 
males, rounded in females. 



SEVENTEEN NEW SPEC1BS of CASTIAR1NA 



49 



Size 

Males, 10,1 =t 0.15 x 3.2 ± 0.05 mm (110). Females, 
107 ± 0.24 x 3.4 =t 0.08 nun (56), 

Aedeagus (Fig. IU) 

Paramcres angled outwards from basal piece, 
rounded at apex. Penis sharp, sides obtusely angled 
away Apophysis of basal piece medium width, .apex 
rounded. Proctiger faintly bilobed, lobes rounded (Fig. 
1 HI). 

Female lermtnalia (Fig, 1 U2J 
Proctiger faintly bilobed, lobes faintly rounded 

Remarks 

C azitrea can be distinguished from all other 
members of the C. paraltela species group except C 
nttnpunctam by the dark blue colour of the elytnt. t 
ociapunciata lias round yellow elytral markings 
whereas they are elongate in C. azitrea. In C azutta 
the aedeagus is shod and broad and m C. odopun> iuia 
it is elongate (Figs IP, IU). The proetigers in both -.exes 
of C. octopunctata have pointed lobes whereas in both 
SBX60 of C azurea (he proctiger lobes are small and 
rounded (Figs 1 PI, I P2, I Ul, I Vl). There appear 
to be a eline in sue within C azitrea. Specimens from 
north of Carnarvon and from the NB wheatbelt areas 
of WA are larger than those fern further easi and south . 
A minority of specimens has the first two yellow marks 
on the elytra fused to form an elongate basal mark. 

Etytnottw 
The species name is derived from F azur. hlue. 



CastwHtta eharientessa sp nnv. 
(FJGS IT. 1 T1, I T2, IK) 

Holorype' cr, 10 km S Dongara, WA, 4 ix 1995, 5. 
Barker, SAMA J 21 307. 

Alhrvpe; 9. 20 km S l.aneelm, WA, 4.x. 1990, 5 
Barker, SAMA I 21 308. 

Pararypes: WA* 2.Cf cZ , Cervantes, 23.ix.l977. M. 
Kt well. MPWA, 2o>a\ 6 9 9, 45 km N Eneabha, 
20.ix.19RO, S Barker & D. I Williams, SAMA; 2 V V , 
200 m N Ledge Pt TO .. Lancelin Rd, 8.x. 1980, S. 
Barker. SAMA. lcr. 29 9, McDcnmd Rock. 
Il.ii.1981. G.J. Keighery, WAMA;2cro\ I 9, Green 
Head, 27.viii.l981, R. P. McMillan, WAMA: 1 a, 1 9 , 
2 km N Badgingarra, M. Powell, 15.ix.l9X4. MPWA. 
l&i Greenough, 26/29.viiiJ9H9 > R. P. McMiilan, 
SAMA; 2ct o\ same data as holotype SAMA; I cr, 
10 km S Dongara, 4ix 1995, S Barker, SAMA. 



Patau? 

Head coppery. Antennae bronze. Pronolum coppery 
Scutellum bronze with blue-green reflections Elytra 
yellow with coppery markings coalesced forming a 
yellow margin two interstices wide; a yellow vuta on 
each side from base to pre-apical area. Ventral surface 
coppery. Legs, femora and tibiae coppery; larsomercs 
bronze- Hairs silver. 

Shape and sculpture 

Head closely punctured, median sulcus present, 
muzzle Short. Antennomeres 1-3 obconie, 441 toothed. 
Pronolum closely punctured, small basal fovea 
extending anteriorly to middle as glabrous line; apical 
margin projecting medially, basal margin almost 
straight, laterally parallel-sided at base, angled 
outwards, rounded 10 widest at middle, rounded and 
narrowed to apex. Scutellum scutiform. glabrous, flat 
Elytra punctate -striate, intervals convex, wrinkled and 
punctured more heavily laterally than medially; 
laterally angled out from base, rounded at humeral 
callus, concave, rounded post-medially and narrowed 
to bispinose apex; both spines minute, margin rounded 
and indented between spinet, apices diverging. Ventral 
surface with shallow punctures, edges ot 'abdominal 
segments glabrous, elsewhere dense, flat, feathery 
hairs. S 7 . males truncate: females rounded. 

Si:r 

Males 12 9 ± 0.:4x,4,3 ± 0.08 mm (12). Females. 
13 5 zt 0.17 x 4J =t 0.13 mm (14) 

.Aedeagus (Fig. I J'l) 

Paramcres angled outwards from basal piece, 
rounded apically. Penis sharp, sides acutely angled 
away, Apophysis of basal piece medium width, rounded 
apically. Proctiger bilobed. lobes near mid-line, blunt. 
jpicil edge straight, rounded later-ally (Fig. I XI). 

Female (erminalia (Fig. 1 T2) 

Proctiger as in male, lobes more pronounced. 

Remarks 

This species can be distinguished from any others 
in the group by the elytra) markings- It is the only 
species occurring in WA which has all Of the yellow 
elytral spots fused to form an elongate yellow vitla on 
each elytron. Some specimens have darker retl 
markings on the elytra than others and these tend 10 
fade ro dark brown in old specimens. Six specimens 
of Ihe type series have a long, separate pre-apical 
yellow mark. Ail specimens except those collected at 
McDerrnid Rocks were taken on the flowers of 
Qiamaelauciuni sp. 

Enmology 

The specific name is derived from Gk charienros, 
beautiful. 



™ 



S. BARKER 



Castiarsna ferruginea sp. nov 
(FIGS IQ. I Ql, 1 Q2. 3H) 

Hohrvpe. o\ Wjalki, WA. I8.ix.l957. S. Barker, 
SAMA ] 21 309. 

Allotype 9 - same data as holotype. SAMA F 21 310. 

Parmvpes: WA. 2 cr 0\ WialkJ, ix.1959, R H Uther 
Baker, WAMA; I Or, *8 km NR Wubio, 17. ix 1970, S. 
Barker, SAMA. 1 C\ Walyahmoning Rock (30 D 38 'S 
IIS 45 E) 9.x. 1972, A Baynes & R. Humphries. 
WAMA 3 r/c, 29 9 , Muekinbudm. 10.x 1979, R. 
P. McMillan. WAMA; 29 9. 8 km E Woolgangic, 
22.x , mo, S. Barker APG Ketnpster, SAMA. £o- 
Southern Cross, x.1981, R. P. McMillan, WAMA; I <* . 
Johnson Lake, 8.xi.l98L D Knowles, MPWA; 3tr cr. 
1 v. Southern Cross, 8.x. 1983. R. P. McMillan, 
WAMA; 2cf o\ Eneabba, 15.x 1985, R. P. McMillan, 
WAMA; lo\ | 9. Dedari. 20x1986, M. Powell. 
MPWA. lo\ 30 km E Lake King, 18. xi 1988, M 
Powell . MPWA . 2 o- -y . ? 9 V . N7T Transmitter. 32 
km E Southern Cross, 21 x 1991, T M. $ Hanlon, 
MHSA; 2cro\ 49 9. Ordari. 21.x 1991. T M > 
Hanlon. MHSA; I 9. Karlgafin. flossy Tolland, 
WAMA. 

Colour 

Head dark coppery with blue-green rcllectmns 
Antennae bronze Pronotnm dark coppery with Wue- 
gi.'cn reflections. Scuiellurn dark purple. Elytra yellow 
with dark maroon markings coalesced Forming a 
liarjOW yellow margin from base to pro -apical area, 
one interstice wide from base, two interstices wide ot 
humeral callus two wide pre-medially continuously 
two wide from post-medially: row of tout yellow spots 
on each side, basal round the remaining three elongate 
Ventral surface coppery Legs; femora and tibiae dark 
copper, tarsi bronze. Hairs silver. 

Shape and sadprure 

Hedd punctured, median sulcus present muzzk 
short Antennomcres 1*3 obcoruc, 4- u toothed 
Pronoturn closely punctured, small basal fovea 
extending anteriorly to middle as glabrous line; apical 
margin projecting medially, basal margin almost 
straight, laterally parallel-sided at base, angled 
outwards then rounded to widest medially, rounded and 
narrowed 10 apex Scutcllum scutiform. elongate, 
glabrous, excavate. Elytra punctate-striate. intervals 
convex, punctured and wrinkled, more so laterally than 
medially; laterally angled out from base, rounded al 
humeral callus, concave, rounded post-medially and 
narrowed to unispinose apex; blunt marginal spine, 
margin indented and straight 10 suture, apices hardly 
diverging. Ventral surface with shallow punctures, 
edges of abdominal segments glabrous, elsewhere with 
dense, flattened, feathery hair. S 7 : males, truncate; 
females rounded. 



Size 

Males, 12.3. ± 0.2 x 4.0 -t 0.07 mm (21). Females. 
13.2 ± 0.22 x 4.3 ± 0.09 mm (15). 

Avdeagus (Fig. IQ) 

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

female termmalia I Fig. I Q2) 

Proctiger bilobed, strongly pointed near mid- 1 inc. 
laterally straight. 

Remarks 

The range of this species overlaps with thai of C 
usndata sp, nov. which is approximately the same 
colour but smaller. They can be distinguished on the 
bH$is of the yellow elytral markings in C fkwNgiftffl 
there are four yellow SpoW on each elyia»n and ifl C 
itsiuloia three, as the first two arc fused forming an 
elongate basal spot and the last two are separate. C. 
femtgtnea has a stngle. large marginal spine at the apex 
ot rhc elytra and C. ustulaia has two small spines The 
aedeagi are different (Figs IQ, IV); both male and 
female proctiger* in both species are strongly lobed. 
those of males arc quite similar in size and shape bm 
prortigers of female C ferruginea arc (note pointed 
than in C UStulata which have black pigment spot> 
at the tip of eavh lobe; these .*pots are absent in the 
other species (Figs I 02. i V2). 

Erytrtfrtngy 

The specific name is derived from L fermpmeus 
rust -coloured 



Castiarina uxtulata sp nnv 
(FIGS IV I VI I V2 3P) 

Hototvpe. c , 8 km E Woolgangie, WA, 22.x 1980, S. 
Barker A P G Kempster, SAMA \ 21 111 

Allotype: 9 , same data as holotype, SAMA I 21 312. 

Pamtypes- WA; I cr, Dumbleyung, 5x1963, H Udell. 
WAMA; lo\ I 9. same data as holotype, SAMA; 
3 9 9. Wialki, 21 ix. 1970. S. Barker, SAMA; 2CfCf, 
9 km SW Walyahmoning Rock, 9.X.1972, A. Bavnes 
& R. Humphries, WAMA; 1 a\ 18.9 km WSW 
Coolgardie. 18.ix 1976. R J. Chinnock, SAMA; I 9. 
Dedan, 8.X.1978, T M. S. Hanlon, WAMA; IC 
Muekinbudm, lO.x.1979, R. P. McMillan, WAMA; 



SEVENTEEN NEW SPECIES Oh CiSl7AJU$A 



si 



1 9, Southern Crass, x.1981, R. R McMillan, WAMA; 
la\ Bullfinch, 2.X.1983, B. Junes, MPWA; I o\ 
Southern Cross, 8.X.1983, R. P. McMillan, WAMA. 
1 9. 28 km NE Peak Charles, 9.x. 1983, G Browning 
& G. Mutze. SAM A. 1 or. I <?, 45 km SW McDermid 
Rock, 24.x. 1985, T. F. Houston & R. W. Thorp. 
WAMA; 29 9, Dcdari, 2G.X.1986, M Powell, 
MPWA; icr, Bindoo Hill Nature Reserve. 27 km W 
Mullewa, 12.ix.1987. T. R Houston, WAMA; 4cr o\ 
29 9. N7T Transmitter, 37 km E Yellow line, 
21.x. 1991, T. M. S. Hanton. MHSA; 2c. 4 9 9- 
Dedari 22. x 1991, T. M. S. Hanlon, MHSA. 3^o\ 
I 9, Karlgann, Bessy Tolland, WAMA 

I oUmr 

Head coppery. Antennae bronze. Pronotum coppery 
Scutellum blue. Elytra yellow with brown markings 
widi coppery reflections coalesced and forming it 
yellow margin from base 10 pre-apical area, one 
interval thick medially, two intervals thick elsewhere, 
ihree medial yellow marks on each side with an 
elongate basal mark termed from the fusion of the basa! 
and pre-medial marks, round post-medial mark and 
elongate pre-apical mark. Ventral surface either all 
coppery or with coppery sternum and coppery-brown 
abdomen with blue reflections. Legs: femora coppery, 
tibiae and tarsi bronze. Hairs silver, 

Shapt* and sculpture 

Head closely punctured, median sulcus present, 
muzzle short. Antennomeres 1-3 obconk, 4-11 toothed. 
Pronotum closely punctured, small basal fovea 
extending anteriorly to middle as impressed line; apical 
maryin projecting medially, basal margin almost 
straight; laterally parallel-sided at base, rounded to 
widest prc-mcdialty. rounded and narrowed to apex. 
Scutellum seutiforrn, glabrous, excavate. Elytra 
punetatc-stnate, intervals convex, punctured and 
wrinkled, more so laterally than medially; laterally 
angled out from base, rounded at humeral callus, 
concave, rounded post-medially and narrowed to 
bispiuose apex; sharp marginal spine, minute sutural 
spine, margin rounded and indested between spines, 
apices hardly diverging. Ventral surface with shallow 
punctures, edges of abdominal segments glabrous, 
elsewhere dense flattened, feathery hairs. S ? ; males 
truncate; females rounded. 

Size 

Males, 121 ± 0.15 x 3.9 =± O05 mm (20). Females, 
12.7. i 0.19 \ 4.0 ± 0.07 mm (18). 

■U^ittigus (Fig. IV) 

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



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

Female wnmnulia (Fig. I V2) 

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

Kermirks 

See remarks under Cfermginea. Elytra! markings 
are the same in this species as in C. antarefiva sp. nov 
but they have non-overlapping ranges. The acdeagus 
in C amurcticd is shorter and broader than that in C 
ustuiaia (Figs. IO. IV). The proctigers of both sexes 
in C. tistulata are bilobed with more highly developed 
lobes than those in C antarctha (Figs I Ol, 1 02. 
I VI. I V2K 

Etymology 

The specific name is derived from L usurious 
scorched 

Casttorina pkaeopus sp. now 
(FIGS IW, I Wl, 1 W2, 3Ej 

Holotxpe: o\ 3 km E Gosnefls, WA, 4.xi.l95b, S 
Barker, SAMA t 21 313. 

Allotype; 9 . Red Hill, WA, lix 1949, R. P. McMillan. 
SAMA I 21 314. 

ftmmyvs; WA: 3o , o\ I v. no data. SAMA; 2Cf or. 
Swan R.. Lea, SAMA; 1 9, Bunbury. W. M. Mack 
i.1898. SAMA; 2 9 9, Penh, xi.1906, SAMA; lacy. 

1 9. Perth, x.1913. SAMA; 4o*Cr, Perth, xi 1920, J. 
W. Mel lor, SAMA: 1 o\ 19, same data as allotype. 
SAMA; I or, Mimtncgarra, Danda/agan. 30 xT955, S 
Barker, SAMA; 2c/ ff, 2 km E Gosnells, 4.xi.V956. 
S. Barker, SAMA; 15 c? c\ same daw as holotypc. 
SAMA. 2crcr, summit Mt Cooke, H\l.xi.l956, S 
Barker, SAMA; \cr. I 9. 70 km SE Perth on Albany 
Hwy, lftxi.1956. S. Barker, SAMA; Icr. foothill* 
Kelmscott, 21.x. 1958, J. Baldwin, SAMA; Jcro\ 

2 9 9, Wilga, 26.x 1972, K. A E Carnaby. SAMA; 
2 a- cr, Lesmurdie, 28. ix. 1955, J. A. Watson, SAMA; 
19, Julimar Forest 24 x 1971, F. H. Uthcr Baker. 
SAMA: 6c?0', 39 9, Cataby Bk, 18x1983, G. 
Browning & G. Mutze, SAMA; Icr, Gosnells. 
7.x. 1980, S. Barker, SAMA; \a, Mundaring Weir. 
3O.ix.1980. T. M. S Hanlon, MHSA; 1 9. Mt Dale, 
29 ix. 1980, T M. S. Hanlon, MHSA. 

Colour 

Head brown with coppery reflections. Antennae 
bronze Pronotum brown with coppery reflection.-.. 



52 



S. BARKER 



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

Shape and sculpture 

Head closely punctured, median sulcus present, 
muzzle short. Antennomeres 1-3 obconic, 4-U toothed. 
Pronottun closely punctured basal fovea extending 
forwards to middle as impressed line: apical margin 
projecting medially, basal margin almost straight, 
laterally parallel-sided at base, angled outwards, 
munded medially at widest part, rounded and narrowed 
to apex. Scutellum scutiform, glabrous, flat Elytra 
punctate- striate, intervals convex, punctured and 
wrinkled laterally, punctured and smooth medially: 
laterally angled oui from base, rounded at humeral 
callus, concave, rounded post-medially, narrowed to 
bispinose apex; very small sharp spines, margin 
rounded and indented between spines, apices hardl> 
diverging. Ventral surface with Shallow punctures, 
edges of abdominal segments glabrous, elsewhere 
dense flattened, feathery hairs. S 7 : males truncate, 
females rounded. 

Size 

Males. II! ± 0.11 x 3.7 ± 0.05 nun (51), K-malcs, 
11.5 ± 35 x $JL ± 0,12 mm (12). 

Aedeagus (Fig. IVV) 

Pararneres angled outwards from basal piece, 
apical ly rounded Penis sharp, sides acutely angled 
away. Hypophysis of basal piece medium width, 
apicully rounded PrftCtfger with small medial notch 
in (ipjcal edge. laterally rounded (Tig- I Wl). 

remole icrnunoini (Fig. I W2) 

Proctigcr bdobed. apical edge straight, laterally 
rounded . 

Remarks 

C piuietfphs sp. oov can be distinguished from all 
others j ( \ ihts complex tfy being the only sptcies which 
bdS the. first three clytral yellow spots Fused to form 
an elongate basal mark with the fourth an elongate pre- 
a -? i c h : r i low matt 

EtymfltK || 

ftv sp'.v.ilc n:mir is derived from Qk pHuitty, 
hi own 



Caxtiarina aniarctica sp. nov. 
(FIGS 10, J Ol, I 02. 3D) 

Holorype: cr , 64 km NF. Esperance. WA. 18.x. W82. 
S. Barker, P. G. Kempster &H. Vanderwoude, SAM A 

1 21 315. 

Allotype: 9 , same data as holotype, SAM A 1 21 316. 

Pamtypes: WA: 1 o% Mt Ragged, 24.x. 1980, S. Barker 
& P. G. Kempster, SAM A; I o\ 13 km N Israelite Bay. 
24.x. |980, S. Barker & P. G. Kempster. SAMA. 1 a. 

2 9 9. 24 km N Israelite Bay, 24 \ 1980, S. Barker 
& P. G- Kempster. SAMA; I o\ same data as holotype. 
SAMA; 59 jr., 7 km N Dempster Kd ScadUcn Rd 
crossing, Esperance district. ISx.1982, S, Barker. P 
G Kempster & H. Vanderwoude. SAMA; 2ao\ 
2 9 9. Parmangocs Rd 2 km NB Clyde Hill TO., 
Esperance district. S. Barker, P G. Kempster & H 
Vanderwoude, SAMA; I 9, Israelite Bay, 21.x. 1982, 
S. Barken P G Kempster & H. Vanderwoude. SAMA 
2 era, 19, 17 km NW Israelite Bay. 21 \.N82. S. 
BarkeT, P G. Kempster & H. Vanderwoude, SAMA 

Colour 

Head coppery Antennae bronze. Pronotum dark 
bronze medially, with coppery reflections laterally 
Scutellum coppery-purple. Elytra yellow with the 
following dark brown markings with coppery 
reflections coalesced leaving yellow margin, two 
intervals wide at apex and at humeral callus, one 
interval wide medially; a row ofyellow spots medially 
on each elytron, basal and pre-medial coalesced 
lorming an elongate mark, post-media! more or less 
round, apical smaller and elongate Ventral surface 
coppery Legs: femora dull purple with coppery 
reflections; tibiae and tarsi bronze. Hairs silver. 

Sluipc and sculpt it re 

Head closely punctured, median sulcus narrow, 
muzzk short. Antennomeres 1 3 obconic, 441 toothed. 
Pmnotum closely punctured, small basal fovea 
extending forward to middle; as glabrous line; apical 
margin projecting medially, basal margin almost 
straight; laterally parallel-sided at base, rounded in 
widest pre-med tally, narrowed to ;ipe>. Scutellunt 
scutiform. tlai. glabrous. HJytra punctate -striate, 
intervals convex, punctured anil wrinkled laterally and 
upically. smooth medially; laterally ;ingle\] oui fie- 1, 
base, rounded at humeral callus, concave, rounded 
post medially and narrowed l<» bispinosc apex; vet} 
small marginal spine, minute medial spine, murcm 
roundel art.J inJcntcd between spines, apices divei^eni 
Ventral surlacc with shallow punctures, etfjjes nl 
abdominal segments glabrous, elsewhere densely ha.iry. 
tiarrs flat and feathery. S maiuti truncate; IvmvjL 
rounded. 



SEVENTEEN NEW SPECIES OF CASTtARIKA 



53 



SIqs. 

Males, 12.0 ± 0,19 x 3.9 A O.Oo mm (14). Females. 
J2.0 ± 0.22 x 4.0 * 0.09 mm (7). 

Aedea$us (Fig. IO) 

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

Female terminalia (Fig. I 02) 

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

Remarks 

The remarks made under those tor C. ustulaia so. 
nov. apply equally to this species as these are the only 
two species in this complex which have this clytral 
pattern they can be easily distinguished by differences 
in aedeagi (Figs LO, IV) and in male and female 
procligers (Figs L Ol, 1 02. 1 VI. I V2>. Also they 
are allopatnc. 

Etymology 
The name is derived from Gk anturktikos, southern 



Cmtiarina nonyma sp. nov. 
(FIGS IX. I XI, I X2, 3A) 

Holotxpe o- . Summit Mt Cooke, WA. 10.xM956, S 
Barker, SAMA I 21 317. 

Allotype; 9 . Juhmar Forest, WA, 24.x 1971, F. H. 
Uther Baker, SAMA 1 21 318. 

Partirypes- WA: ley o\ 2 9 $. Beverley, E K du 
Boulay. SAMA. 2a a , Perth, SAMA: 1 a\ Swan H.. 
SAMA, I 9. xii.1913, SAMA; Icycf, same data as 
holotvpe, SAMA; la, Mt Walker (32*05' S 118*45' 
F) 16 x. 1979, R. P. McMillan, WAMA, I o\ GoMiells, 
7*1980. S. Barker, SAMA. 19. Fonnsi field, 
27.viii.l978, T M. S. Hanlon. WAMA: 1 9 . EfleObta, 
17.x. W85. R P McMillan, WAMA. 

Colour 

Head and antennae dark maroon. Pronotum dark 
maroon with btue reflections medially. Scutettum dark 
maroon. Elytra yellow with maroon markings 
coalesced to form yellow margins, two intervals wide 
at humeral callus and apically. one interval wide 
medially ; a medial row of tour yellow spots on each 
elytmn, basal and post-mediaJ more or less round prc- 
niedial and pie-apical elongate, in about a quarter 09 the 



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

Shape and sculpture 

Head closely punctured, median sulcus shallow, 
muzzle short. Antennonieres 1-3 obconic, 4-J1 toothed. 
Pronotum closely punctured, small basal fovea 
extending forwards to middle as impressed line; apical 
margin projecting medially, basal margin almost 
straight; laterally parallel-sided at base, rounded to 
widest medially, rounded and narrowed to apex 
Scutellum scutiform, glabrous, excavate Elytra 
punctate-striate, intervals convex punctured and 
wrinkled laterally and apically. smooth medially, 
laterally angled out from base, rounded at humeral 
callus, concave, rounded post-medially and narrowed 
to unispinose apex; small, blunt marginal spine, margin 
straight and indented to suture, apices diverging. 
Ventral surface with shallow punctures, edges of 
abdominal segments glabrous, elsewhere with dense 



hair. S-,'- males* truncate; females 



flat, fealhery 
rounded. 

Sre 

Males, 11.0 ± 0.28 x 3.5 ± Oife mm (lfiX Females, 
111 ±. 0.24 x.V7 ± 0.07 mm (6), 

Aedea^us (Fig IX) 

ftmuneres angled outwards from basal piece, slightly 
rounded post-medially then angled outwards, apically 
rounded. Penis sharp, sides acutely angled away. 
Hypophysis of basal piece medium width, apically 
rounded. Proctiger with medial apical edgjc straight. 
then angled forming I wo small broadly pointed lobes, 
laterally rounded (Fig- I XI), 

Female lermmalia (Fig. 1 X2) 

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

Remarks 

No female specimen*-, associated with males at the 
*ame collection locality were available This and the 
following species C eni^mu sp. nov have elytra 
predomtnently eight spotted, although a small number 
of each has the first two spots coalesced The two 
species can be distinguished by differences in acdea^u 
which are short and broad in C enigttui and elongate 
in C". nonyma (Figs \K, IV) and in male and female 
piocbgers which arc virtually unsculpturcd in C 
enigma and bilobed with pointed lobes in C nonynu: 
■Tigs ! XI, I X2, 1 Yl. I Y2). The distribution of t"- 
itonvma appears to be mainly to (he cast of the Darling 
Scarp fault line, while that of C enigma is to the west 
of the Darling Scarp on the coastal Plain, 



54 



S. BARKER 



Etymology away. Apophysis of basal piece medium width, apically 

The name is derived from Ok anonymos, unknown rounded. Proctiger broadly rounded al apex, sides 

rounded (Fig. 1 Yl). 
Castiarina enigma sp. nov. 

(FIGS 1Y, 1 Yl. 1 Y2, 3C) Female terminalia (Fig. I Y2) 

Holotype. OS Regans Ford. WA, 9 x.1970. K & E. ^^ l '° lM 

Carnaby. SAMA I 21 319. $m ^ 

Allotype: 9. 6 km S Gin Gin, WA, 30.ix.l956. S. The remarks under C nonyma apply equally to this 

Barker. SAMA ] 21 320. species. 



Paratypes: WA 1 or, 19, no data, SAMA. I 0>, E 
Ashby. SAMA; I 9. Perm. SAMA. 1 9 , feUi .vi 1905 

SAMA; 2<7 0-, Perth, x.1913, SAMA; 2o-o\ 19, 
same dala as allotype, SAMA. Her cr. 1 Q , same Jala 
as holotype. SAMA. 

Colour 

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

Shape and sculpture 

Head closely punctured, median sulcus shallow, 
muzzle short Antennomcres 1-3 obconic, 4-11 toothed 
Pronotum closely punctured, narrow basal fovea 
extending forwards to middle as glabrous line; apical 
margin projecting medially, basal margin rounded from 
base to widest medially, rounded and narrowed 10 ape*. 
Scutellum scutitorm, glabrous, Hat. Elytra punctate- 
striate, intervals convex, punctured and wrinkled 
laterally smooth medially; laterally angled out from 
base, rounded at humeral callus, concave, rounded 
post* medially and narrowed to bispinose apex; small. 
blunt marginal spine, minute sntural spine, margin 
rounded and indented between spines, apices diverging. 
Ventral surface with shallow punctures, edges ol 
abdominal segmcnLs glabnnus, elsewhere densely hairy, 
hairs flat and leathery. S 7 : males truncate, females 
munded, 

She 

Males. 10.7 ± 0.17 \ 3.4 ± 0.06 mm (15). Females. 
U.8 ± 0.37 x 4.0 -± 0.13 mm (6) 

Aetleai>us (Fig. 1Y) 

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



Etvmoh^y 
The name is derived from I. aenigma. mystery 



Castiarina aura sp. nov. 
(FIGS IS. I SI. I S2 ; 31) 

Holotype: o\ 131 kmSExrnouth, WA. L2.tx,l984. M 
Powell, WAMA. 

Allotype: £ , same data as holotype. WAMA. 

Paratypes: WA. 19, 50 km N Kalbam TO., 
20.viii.1978, T. M.S. Hanlon. WAMA; I o\ YirdicCk, 
I8.vni.l983, M. Powell. MPWA. I cr Coral Bay. 
10,ix.»984, M. Powell. MPWA; IO\ 1 9, Carnarvon, 
28.vtii.l987, A. Hay. SAMA; J Q, 94 km S Learmonth. 
2.ix.l995, Powell & Kershaw; MPWA; I 9, 62 km S 
Leawionth, 4.ix.l995. MPWA; Io\ I 9, 26 km S 
Learmonth, 3 ix.|995. Powell & Kershaw, MPWA. 

Colour 

Head coppery. Antennae bronze with coppery 
reflections. Pronotum coppery, with medial blue-green 
reflections. Scutellum blue-green. Elytra yellow with 
the following elytra! markings: markings coalesced, 
coppery apically, with blue-green reflections along the 
suture and ewer ihe humeral callus forming a yellow 
margin two intervals wide, medial row ol four yellow 
spots on each elytron, basal and post-medial more oi 
less round, pre-medial and pre-apical elongate. Ventral 
surface and legs coppery. Hairs silver. 

Shape and sculpture 

Head closely punctured, median sulcus shallow, 
muzzle short. Antennomeres 1-3 obconic, 4-11 toothed. 
Pronotum closely punctured, basal fovea extending 
forwards to middle as glabruus line; apical margin 
projecting medially, basal margin almost straight, 
laterally parallel-sided at base, munded to widesi before 
middle, rounded and narrowed to apex. Scutellum 
sculifonn, wrinkled, excavate. Elytra puncuttc striate- 
intervals convex, punctured and wrinkled laterally 
smooth medially; laterally angled outwards from base 



SEVENTBEN NEW SPECIES OF C4SI7AR1NA 



55 



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

Size 

Male, 11.9 ± 0,42 x 4.0 ± 0.15 mm (5). Females, 
12.9 ± 0,44 x 4.3 ± 0.18 mm (6). 

.Aedeagus (Fig. IS) 

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

Female terminalia (Fig. 1 S2) 

Procliger bilobed, lobes strongly pointed near 
mid-line. 

Remarks 

C aura sp. nov, and C ferruginea sp. nov. arc 
similar in that both have four separate yellow spots on 
each elytron and a large single marginal spine on the 
apices of the elytra although the elytra! colour is 
different, C. aura being red with green reflections 
while C. ferruginea is brownish. Aedeagi differ as they 
are shorter and narrower in C. aura than they are in 
C. ferruginea (Figs 1Q, IS). The proctigers of C 
ferruginea males are strongly bilobed while those of 
C aura are faintly bilobed (Figs I Ql. I SI). The 
proctigers of females of both species are bilobed and 
pointed but the lobes arc further apart in C aura man 
they are in C. ferruginea {Figs 1 Q2, 1 S2). 

Etymology 
The name is derived from L aura, glow. 

Key 10 W\ species of C parallela complex 

!. Elvira buck ground colour dark blue. 1 

Llytfii background uiluur hmwn. 

red or green 3 

2. Elytra with 8 round, yellow mark* 



Hlytra wilJi 2 round. 6 elnngaie 

yellow murk* 

Elvira bright rod or partially 

or wholly brassy green 

Hlytra brown or red-brown 

Elytra bright red, elyiral apices with two 

small spines, elytra) spots coalesced into 

single elongate yellow mark 

on each side 

Elytra bright red or partially or whollv 

brassy green, elytra with H yellow spots 



nrioftutiiltttu 
Barker 



arurcu sp, nov 



(hiirimcssd sp, nov 



Elytral apices with 1 large spine 


aura sp. nov. 


Elytral apices with 2 small, 




blunt spines 


crochvhr EL & G) 


Some elytral marks coalesced forming 




6 or fewer yellow marks 


7 


Hlytra with 8 yellow marks 


10 


First 3 yellow marks on each elytron 




coalesced, forming 1 elongate anterior 




mark and 1 small elongate mark 




posteriorly on each side 


pharopus sp. nov. 


First 2 yellow marks on each elytron 




coalesced, forming 1 elongate anterior 




mark and 2 small elongate marks 




posteriorly on each side 


i 


Pronotum bronze, elytra dull hmwn. 




nale procliger slightly sculptured, 




female proctiger unsculptured. Largest 




member Of group. 


adttsia sp nov. 


Pronotum. elytra with coppery reflections 


9 


Proctiger bilobed, lobes pointed, in 




females with pigment spot ai up 


ustulata sp. nov, 


Procliger medially notched, lobes blunt 




wiihoui pigment in females 


antaraica sp, nov. 


Elytral apices with single large spine 


ftrru\>ttu-u *p nOV. 


Elytral apices bispinose 


11 


Aedeagus broad, procliger unsculptured 




in both sexes 


t'niamu sp. nov, 


Aedeagus narrow, proctiger sculptured 




in both sexes 


nonyrtut sp. nov_ 



Castiarina nullarborica sp. 
(FIGS U, 4D) 



nov. 



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

Allotype 9 , Nullarbor Plain, SA, SAMA I 21 381, 




,, r , i mm. mm 



Fig. 4. Habitus illustrations of the following Cusliurina 

arocjey A. Castiarina jaekfutsenpuschi sp, nov. holotype. 
B. C hemhhstem sp. nov. holotype C. C. paullutsenpusehi 
.sp. nov. holotype. D. C*. nttllarbunctt sp. nov. holotype. 
h. C. tlettutnj sp. nov. allotype. 



56 



S. BARKER 



Pamrypes: SA: lo\ 32 krn E HucJa, Jl.xii.1984, M. 
Powell, MPWA. WA; I o*. same data as holotypc, 
SAMA 

Colour 

Head dark blue. Antennae blue-green. Pronotum 
hronze medially, dark blue laterally. Scutellum dark 
blue Elytra yellow with red margin and die following 
black markings with blue reflections: narrow basal 
margin- broad pre-medial fascia nol reaching margin, 
distally angled anteriorly: broad post-medial Ium-i.i 
reaching margin, spade-shaped apical mark covering 
.ipex and spines, last iwo marks connected broadly 
along suture. Ventral surface bronze. Legs dark bluc- 
Hairs silver, 

Shape and sculpture 

Head closely punctured, median suJcus broad, 
muzzle short. Antennomcres Mobconie, 5-11 toothed 
Pronotum closely punctured, small basal fovea; api&l 
margin straight, basal margin bismunte; laterally 
parallel-sided at base, rounded to widest prc-medially, 
rounded and narrowed la apex Scutellum scutiform. 
glabrous, flat. Elytra punciate-stnate. intervals convex, 
puncrured; laterally angled out from base, rounded al 
humeral callus, concave, rounded post-medially and 
narrowed lo bispinose apex, sharp marginal spine, 
small, sharp suturaJ spine margin rounded and deeplv 
indented between spines, apical margin subserrate, 
Ventral surface with shallow punctures, edges of 
abdominal segments glabrous, elsewhere moderately 
hairy, hairs medium length. S 7 : truncate in both sexes. 
Male legs 2-3, pulvilli absent on tarsomeres 1-3 
replaced with a small double, median spine. 

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

Aedeogus (Fig U> 
Wedge-shaped. 

Remarks 

This species is closest to the morph of C plaa'da 
(Thomson) which has a red margin and occurs On the 
west coast of WA and on Rottnest Is.. WA. C. 
nullarborica is a smaller species than C. placida and 
(he male genitalia differ in size and shape (Figs II, U» 

Etymology 

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



Castiahna demarzi sp. (V-^. 
(FIGS IH, 4E) 

Hotorxpe: o\ Eurardy HS (27°34'S, 114*40 'E) WA 
I9.vh1.1980, C A. Howard & T. F. Houston. WAMA 



Allotype. <?, 36 km NE Tamala Station, Shark Bay 
WA.28.ix. 1988, D. Knowlcs. WAMA. 

Paratvpes; WA 1 9, same data as allotype, MPWA; 
1 9, 26 km NE Tamala Station, 6.x.l'J88, D. Knov*. l.s 
SAMA: 1 9 , found in seed collection from NW coast. 
29.viii.l986, H. Dcinarz, HDWA. 

Colour 

Head, antennae and pronotum bronze with or 
without coppery reflections. Scutellum blue or bronze 
I Ivira yellow with the following black markings with 
blue reflections, narrow basal margin; pre-mcjf.il 
fascia not reaching margin, distally angled anteriorly, 
post-medial fascia reaching margin, projecting 
anteriorly in middle of each elytron; mark covering 
whole apex, marks connected along suture in holotypc 
but not allotype. Ventral surface and legs coppery - 
Hairs silver. 

Shape ami sculpture 

Head closely punctured, median sulcus broad, 
muzzle very short. Antcnnomeres compressed, 1-3 
-•huTik-, 4-1! toothed, Pronotum closely punctured. 
basal fovea extending anteriorly to middle as glabrous 
line, b;isal notches represented by glabrous area on each 
side closer to margin than middle; apical margin 
projecting medially, basal margin barely Insinuate; 
laterally angled outwards from base, rounded to widest 
after middle, rounded and narrowed to apex. Scutellum 
scutiform. glabrous, Hat. Elytra punctate-striate, 
intervals convex, punctured: laterally angled oui from 
base, rounded at humeral callus, concave, rounded 
post-medially and narrowed to bispinose apes; sharp 
marginal spine, small* sharp sutural spine, apices 
diverging, apical margin subserrate. Ventral surface 
With shallow punctures, edges of abdominal segments 
glabrous, elsewhere short sparse bait S,: maK- 
truncate'. females rounded. 

Stzc 

Male. 10,0 x 3.7 mm (1). Females, li.7 =t 0.22 x 4.f> 
i 0.11 mm 14). 

Aedeugus (Fig. IH) 

Parameres parallel-sided from basal piece, rounded 
prc-medially then narrowed to apex. Penis blunt, sides 
acutely angled away. Hypophysis tit basal piece 
medium width, apically rounded. 

Remarks 

The structure and clytral markings of this species 
resemble C. hueolica (Kerremans). However C 
hueolica has head, pronomm and ventral surface green 
and there are size- differences between aedeagi, thai 
of C demuni being smaller man that of C bucoiua 
(Figs |G, IH) 



Etymology 

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



Castiahna jackkastnpusthi sp. nov. 
(FIGS 1L. 4A) 



SEVENTEEN WW SPECIES OF CAS71ARiHi 

Remarks 



P 



Holotypt'- tf . Card well Ra,. 
NSGOpuStih. SAMA I 21 M2. 

Allotype: v, C&rdwcll Ra,. 
Hasenpusch. SAMA I 21 323. 



Old. 22.Klijy§5, J 



Qld, 22.xii.W9S, r 



Colour 

Head reddish-bronze, muzzle green-bronze. 
Antennae green Pronotum reddish-bronze with a 
curved blue bar. concave inwards, on each side Of the 
mid line from base |p apex, Scutellum green with 
yellow reflections. Elytra yellow with the following 
black markings: broad basal margin: broad pre-medial 
fascia with ends expanded anteriorly reaching basal 
margin and posteriorly reaching margin enclosing a 
yellow basal spot and a yellow spot on margin at 
humeral callus; broad post-medial fascia reaching 
margin" and mark covering whole apex; yellow medial 
lascia nol reaching margin; yellow post-medial fascia 
tTOl reaching suture or margin; ventral surface green 
with yellow reflections, legs blue-green. Hairs silver. 

Shape and sculpture 

Head closely punctured, median sulcus present, 
muzzle short, Antennomeres 1-3 obconic, 4 half- 
toothed, 5-11 toothed. Pronotum closely punctured, 
small basal fovea; apical margin straight, basal margin 
bisinuate; laterally angled inwards from base, rounded, 
widest before middle, rounded and narrowed to apex. 
Scutellum seutiforrn, punctured, flat, Elytra punctate- 
striate, intervals convex, punctured; laterally angled 
out from base, rounded at humeral callus, concave, 
rounded post-medially, narrowed to bispinose apex, 
large sharp marginal spine, minute sutural spine, 
margin indented and straight between spines, apices 
hardly diverging. Ventral surface with shallow 
punctures, edges of abdominal segments glabrous, 
elsewhere sparse very short hairs. S 7 : male rounded: 
female slightly rounded and turned under. 

Male, 6.4 x 2,5 mm (I) Female. 6.9 x 2.6 mm li). 

.Aedeagus (Fig- 1L) 

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



This species superficially resembles C. ndisTa 
(Rainhow). (t is however, smaller, the structure of the 
antennomeres differs as the 4th antennomere of C. 
cvdtsui is fully toothed and male genitalia are dissimilar 
(Fags IK, 1U. 

Etymology 

This species is named to honour Mr J Hascnpusch. 

Innisfait, who has generously supported my research 
by loaning specimens and providing information. 



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

Holotype: o\ Marsupial Ck iicut Ctoydoii. Old, 
2vi.l995. P. Hasenpusch. SAMA I 21 334. 

Allotype: Q , same data as hoiotypc, SAMA I 21 325 

Paratypes: Qld: 1 9. Saiflfi data as hoiotypc JHQA. 
2 era. Marsupial Ck, 1-15 iv.N95, P Hasenpusch, 
JHQA; lor. 1 9. Marsupial Ck, 2.vi.I995. J. 

Hasenpusch, JHQA. 

Colour 

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

Shape and sculpture 

Head closely punctured, glabrous, median sulcus 
present, muzzle very short. Antennomeres 
compressed. 1-3 obconic, 4-11 toothed. Pronotum 
closely punctured, glabrous, basal fovea extending 
forwards but not reaching middle, basal notches on 
each side closer to margin than middle; apical margin 
projecting medially, basal margin bisinuate; laterally 
parallel-sided at base, rounded to widest at middle, 
rounded to apex. Scutellum seutiforrn. glabrous, 
excavate. Elytra punctate-striate, intervals convex, more 
so apically; laterally angled out from base, rounded 
al humeral callus, concave, rounded post-medially, 
narrowed to bispinose apex; shaqi marginal spine, 
small sharp sutural spine, margin indented and rounded 
between spines, apices diverging, apical margin 
strongly sub-serrate. Ventral surface with shallow 
punctures, edges of abdominal segments glabrous. 



58 



S. BARKER 



elsewhere moderately hairy, hairs medium length. S r 
males truncate; females bilobed, each tobc with four 
claws. 

Size 

Males, 13.2 ± 0.16 x 4 5 ± 0.07 mm 14). Females. 
13.5 i 0.45 \ 4,7 =t 0.18 mm (3) 

Aedeagus (Fig. IF) 

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

Remarks 

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

Etymology 

The species name honours Master Paul Hasenpuseh 
its discoverer. 



Castmrina hemizostera sp. nov. 
(FTGS IN, 4B) 



Halotype: o\ Cardvvcll 
Hasenpuseh, SAM A 1 21 



Ra.. Qld. 22 Nii im .1 

326. 



Allotype: Q , Cardwell Ra., Old, 24.xii.l993, J. 
Hasenpuseh, SAMA 1 21 327. 

Paratopes: Qld; I o\ Cardwell Ra-, 19.xii.1995, J. 
Hasenpuseh, JHQA; 2c o* , same data as holotype. 
JHQA; 1 o\ 1 V- 22.xii.l995. P. Hasenpuseh, JHQA, 
3c? cr, 3D.Xil 1995, J. Hasenpuseh, JHQA. 

Head black with blue-green reflections, muzzle blue. 
Antennae green. Pronolum with purple-green 
reflections medially, blue-green laterally. Scutellum 
green. Elytra yellow; with black markings with blur 
grren and/or purple rcfleetinns coalesced leaving the 
following yellow marks; pre-medial yellnw fascia 
reaching margin bui not suture, broad pre apical vellum 
la m i.i reaching margin hut not suture. Ventral surlace 
black with bronze reflections. Legs blue. Hairs silver 



Shape and sculpture 

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

Size 

Males, 7.0 ± 0.2 x 2.5 * 0.09 mm (8). Females. 
7.9 x 3.0 mm (2). 

Aedeagus (Fig. IN) 
Parameres angled 



outwards from basal piece. 



rarameres ungicu nmwaius nvra uoaai pic*.*.. 
>unded and widened post-medially, rounded at apex 
enis sharp, sides acutely angled away. Apophysis ot 



ro 

Penis sharp, ^i^ H w UM * v 

basal piece wide, apically rounded 

Renuirks 

This species is allied to C. bella (Saunders) and is 
closest to C. aglaia (Barker). However, the posi-rnediat 
fascia in C aglaia is red and the male genitalia differ. 
(figS IM, IN) 

Etymology 

The name is derived from Ok hemisys, half, Gk 
zoster, bell. 



Acknowledgments 

I nm endebted to the following for assistance Dr 
T R Houston, WAMA; Dr K. Walker, NMVA, Mr 
T, Weir, AN1C; DrE. G. Matthews, SAMA, Mr D 
.1. Williams, Mr P. G. Kempster and Ms H 
Vanderwoude, Department of /oology. University of 
Adelaide. I am endebted CO the following collectors hir 
the loan and gift of specimens; Dr. F H Utbei Bakei, 
Cottesloe; Mr H. Demarz, Guildenon; MrT M. S 
H anion. Hunters Hill; Mr J. Hasenpuseh. Innisfail: Mr 
R. P. McMillan, Kallaroo; Mr M Powell, Melville. 
Mr S. Watkins. Caparra. 



References 



Aknl-ii R. II. Jr. &VMUELSOM C A- &- NistutM, G. M 
il'Wl The Insect and Spider collections nf the world" 
2nd cd (Sandhill Crane Press, Gainsvilk i 

Bakkj*. S. (WK7> Fighteen new species of Stiy t modera 



iCasfiarina) (Coleupiera: Buprcstidae). Trims. R. Soc< V 
AUSL Ul, 133-146. 

(»W*> Seventeen new species of Australian 



Rupreslidae (Insecta: Colcoptera) and a host plant m 
Citstiarina ttptmii (Barker). fhiJ. 117 \5 2t> 



SEVENTEEN NEW SPECIES OF CASTIARINA 59 

(1995) Eight new species of Australian Buprestidae Kerremans, C. (1890) Especes inedites du genre Stigmodera 

(Insecta: Coleoptera). Ibid. 119, 149-156. Eschscholtz. Bull. Soc. Ent. Belg. 1890, 40-49. 

Blackburn, T. (1890) Further notes on Australian Laporte, F. L. & Gory, H. (1838) "Histoire naturelle et 

Coleoptera, with descriptions of new genera and species, iconographie des insectes coleopteres, publiee par 

Ibid. 13, 121-160 monographies separees" Vol 2 Suite aux Buprestidae (P. 

Deuquet, C. M. (1956) Notes on Australian Buprestidae, Dumenil, Paris). 

with descriptions of three new species and two subspecies Saunders, E. (1868) A revision of the Australian Buprestidae 

of the genus Stigmodera, Subgenus Castiarina. Proc. Linn. described by the Rev. F. W. Hope. Trans. Roy. entomol. 

Soc. N.S.W. 81, 153-156. Soc. Land. 6, 1-67. 

Gehin, J. J. B. (1855) Coleopteres nouveaux ou peu Connus Watt, J. C. (1979) Abbreviations for entomological 

Decade 1. Bull Soc. Hist. nat. Men. 7, 53-65. collections. N.Z. Zool. 6, 519-520. 



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

IN AUSTRALIA 



By Peter Kolesik* 



Summary 

Kolesik, P. (1996) A new genus and three new species of Cecidomyiidae (Diptera) 

from Olearia spp. (Asteraceae) in Australia. Trans. R. Soc. S. Aust. 120(2), 61-67, 31 

May, 1996. 

A new gall midge genus, Trigonomyia, and three new species, T. ananas from Olearia 

ramulosa (Labill.) Benth., T. cristata and T. tulipa both from O. axillaris (DC.) F. 

Muell. Ex Benth., are described. Detailed descriptions of the adults, larvae, pupae and 

galls are given. The species are distinguished from each other by both their 

morphology and the appearance of their galls. The new genus is diagnosed and placed 

in the tribe Oligotrophini within the supertribe Lasiopteridi of the subfamily 

Cecidomyiinae. 

Key Words: Cecidomyiidae, Trigonomyia ananas sp. now, Trigonomyia cristata sp. 

nov. , Trigonomyia tulipa sp. nov. , Olearia ramulosa, Olearia axillaris, South 

Australia. 



Ihmsaciitms of the Hoval Society <>JS. Aust, (W96J* 120(2). 61-67 

A NEW GENUS AND THREE NEW SPECIES OF CECIDOMYIIDAE (D1PTERA) FROM 
OLEAR1A SPP. (ASTERACEAE) IN AUSTRALIA 

by Peter K0LESK* 



Summary 

Kolesik.. P. H996) A new genus and three new species. Of Cecidomyiidae (Dipiera) from Ulearia spp. (Asteraceae) 
in Australia, Trans- R S»c. S. Aust. 120(2), 61-67. 31 May. 1996. 

A new gall midge genus. Tri%onomyia. and three new species, T, unarms from Oleariu rumulo.sa (Labfll.) Bcnth., 
T iTistata and T. tutipti both from O. axillaris (DC ) F. Mudl. ex Benin,, are described Deluded descriptions 
of the adults, larvae, pupae and galls arc given Tne species are distinguished from each other by both their 
morphology and the appearance of their galls. The new genus is diagnosed and placed in the tribe OligotTophini 
within the supenribe Laskipieridi of the subfamily Cecidomyiinac. 



Key Words; Cecidomyhdae, Trigonomyia ananas sp. nov., Trigowwyia enstatu sp. nov. 
sp. nov., Olearia ramulosa, Oleuria axillaris. South Australia. 



Tritfttrutmyia tulifta 



Introduction 

Three new gall midge species arc described here that 
were found galling flowers of two species of the daisy- 
bush, Oleariu Moench (Asteraccae). Triganomyia 
ananas sp. nov. was found in Black Hill Conservation 
T A ark, near Adelaide, infesting the twiggy daisy-bush, 
(). ramulosa (Labiil.) Benth. Tngcmomyia crktaw sp 
nov and T. tulipa sp. nov. were discovered at 
Beachport, in the Lower South-East of South Australia, 
both attacking the coastal daisy-bush, O. axillaris 
<D.C.) P. Muell. ex Benth. 

Olearia includes some 75 species in Australia and 
25 in New Zealand and New Guinea (Cooke 1986). 
Olearia ramulosa is an aromatic shrub, about 1.5 
metres high, much -branched, with a woody stem and 
numerous, small, yellow-white flowers which occurs 
ihroughoul Australia in mailee, woodland and coastal 
scrub (Cooke 1986). It is common in Black Hill 
Conservation E^rk where it often forms dense localised 
populations on poor stony soils. Olearia axittaris is 
a 2-3 metres high shrub, morphologically distinguished 
from (I ramulosa by larger leaves and minute ligules. 
Olearia axillaris forms a dense scrub on coastal sand 
dunes of moderate and temperate Australia (Cooke 
1986) and is a dominant plant along the Beachport sea 
shore. 

A new genus is proposed tor the three new gall 
midge species, tt is placed in the subfamily Ceeido- 
myiinae and supertribe Lasioptcridi. Tt is compared 
to RhapaUtmyia of the tribe Oligotrophia from which it 



* Department of Horticulture, Viticulture and Oenology 
University of Adelaide PMB I Glen Osmond S. Aust. 5064 



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

Material and Methods 

Three- distinct kinds of flower galls were sampled 
One was collected from O. ramuhsa in Black Hill 
Conservation Park near Adelaide (I7,ix.l994) and two 
from O. axillaris on coastal sand dunes at Beachpor. 
(6.X-1994) The two types of galls collected from O. 
axillaris were kept in separate bags and all galls weir 
processed in two ways according to the method 
previously described (Kolesik 1995). Microscope 
mounts of the type specimens were prepared by 
maceration in 20% KOH, followed by processing 
through distilled water, 70% and 99% ethanol, xylene 
and were mounted in Canada balsam for examination 
by phase-contrast and bright-field microscopy. Larvae. 
pupae and pupal skins were mounted dorso-ventrally 
Adults were dissected into four (females) or five 
(males) pieces and their particular parts mounted 
separately: wing, head froniaily. thorax laterally, female 
abdomen dorso-ventrally or laterally and male genitalia 
and abdomen dorso-ventrally. Measurements were 
made with an eyepiece graticule. Drawings were done 
with the aid of a camera lucida. The type series and 
other materials retained in 70% ethanol are deposited 
in the. South Australian Museum, Adelaide [SAM), the 
Australian National Insect Collection, Canberra 
|AN1C] and the United States National Museum, 
Washington DC [USNM). Adult terminology follows 
usage in Gagne (1981) Both larval and pupal 
terminology follows Gagne (1994). 



t>: 



P. KOIXS1K 



Genus Tri$onomykt gen. nw, 

Type species. THfiammyia ananas 
■■\iiuh\ 

Wiu^s v*itll R> joining C ai win/, apex, R- absent 
Ri jninirvjj f* near wing mid-length, M* absent, Ofl 
forked. Maxillary palpus with 3 segments, palpiger 
well developed. Eye facets rounded, eye bridge 2-4 
beds medially. Anlenna with variable number ot 
fhigellomcrcs. usually 16-18. first and second only 
weakly separated Fla^eilomeres cylindrical with neck 
longer in male than in female, with long and .stout setae 
ui up lo three whorls; circnmfiUr kxirw short . forming 
sparNe network, similar in Kith sexes Empodia longer 
than claws, pulvilh stout, about half daw length. Claws 
simple, broadly curved. Abdomen: tejviie I scleroti/ed 
iti hoth sexes, with posterior sctaJ row only, tcigitcs 
U - VIII in male and It - VII in female seteroii/ed, 
with single posterior scial row interrupted mesally. pair 
rtf sparse scial fields laterally and one seta in both 
anterior corners, female tergite VII] not sclcrotizcd, 
with triangular field of scattered setae at posterior end; 
stcrmies II • VIU in male and II - VJT in female 
sclcrotizcd, with dense, uninterrupted posterior band 
o| seiae. scattered setae anteriorly and isolated pair of 
setae on posterior end. Male genitalia: ^otiocoxjtes 
cylindrical, uidobed, setose and setulose; gonosiylus 
situated caudally on gorioeoxite, cylindrvcaK slightly 
tapenng towards ape*, with short apical tooth 
comprismg one claw and several firm bristles, setose, 
setulose throughout; cerci bilobed, with several s?out 
setae on each lobe, setulose; hypoproet bilobed, with 
one long sew on each lobe, setulose; parameres divided 
into two parts, basal lobe simple, asetose. setulo&e. 
apical lobe asetulose ? bearing 5-6 parallel running 
lamellae, asetulose, bearing altogether 6-8 larpe, setose 
papillae; aedeagus robust, strongly sclcrotizcd vcnrtrally 
and npically, with apical cud triangular. Ovipositor: 
prolnisible; cerci fused into single, terminal lamella. 
macular in dorso-ventral view., with numerous strong 
setae, setulose; hypoproet trape/okl in dono-vcmtal 
view, short, bearing 2 setae posteriotty, seiulo»e. 
Fupa 

Integument of abdominal segments covered by 
spiculae. Prothorax and abdominal segments t-VUI 
wilh spiracles. Antenna! horns short, angular. Cephalic 
pair of papdlae with strong, long setae. Frons with one 
pair of upper frontal weakly sclcrotizcd depressions 
and one ot* 2 lower facial papillae on each side with 
short seta. Abdominal segments I-VII with 1 pair of 
ventral papillae, 2 pairs of pleural papillae aixl 3 pairs 
ol dorsal papillae. abdominal segments VIU and IX 
with I pair of ventral, 2 pairs of pleural papillae and 
I pair of dorsal papillae. All papillae setose. 
Utno 



Head strongly sclerotized, posterolateral apoderncs 
shorter than one fourth of head capsule lengm, 
antennae about three times longer than wide at base 
.ontcal. Neck segment with I pair of setose pleural 
papillae Thoracic segments without spatula but wilh 
depression where spatula would normally .appear. I 
pair of ventral papillae, 2 pairs of lateral papillae. -■ 
pairs of pleural papillae, 3 pairs of dorsal papillae 
Abdominal segments 1-VII with two sternal 
depressions, I pair of ventral papillae. 2 pairs ol pleural 
papillae, 3 paiis ol dorsal papillae. Abdominal segment 

VIII with 2 pairs of ventral papillae, 2 pairs of pleural 
papillae, I pair of dorsal papillae. Abdominal segment 

IX with 4 pajrs of terminal papillae. All papillae setose 
except thoracic lateral ones. Setae long and clearly 
apparent on all papillae with exception of ventral 
papillae which are only slightly longer than iniegU- 
rnental spiculae. 

TrifOftfWffa combines "tugomin; Ok lor triangle 
winch reters in the <hapc nl the apical end of aedeagus 
and "myia" Gk for fly. commonly used as sutfut for 
genera of Ccodomyildac. 

Remarks 

Trignnomyia belongs to the tribe l.asiopterid) because 
it has male parameres and an irregular number of 
antenna! flagcllomeres. The new genus belongs to the 
OJtgolrophim ami within that tribe to a group of genera 
that includes Rhopalomyia . and for which the tribal 
name Oligotrophia (in the strict sense, not including 
Dasineurini), is available. These genera share the 
tallowing derived characters, reduction of the palpus 
to three or fewer segments, relatively short parameres 
dial do not clasp the aedeagus along its full length, 
larvae each living in a separate cell in galls with 
pupation occurring inside the larval cell. Other, 
probably primitive characters shared by all members 
of Rhopahmyla and relatives are the completely 
wtulose male gonostylus and the never divided female 
eighth abdominal tergite. even when the ovipositor is 
elongate (Gagne* et ai in press) - Trigntwmyia differs 
from Rhopahmyia in several ways. In Trigonomyia the 
parameres arc divided into two distinct lobes, ihc 
aedeagus is sclerotized apieally and the larva has setose- 
collar papillae. None of these characters has been noted 
in Rhapalamyia or related genera, although presumably 
separately derived, indistint to distinct divisions can 
be found in die parameres in Dmweurt} is L). 
tMinptrra and Lfdomyio (Gagnc 1994}. 

Trigcnomyia ananas sp. nov. 
<F1GS I, 2. 5-8, 11-17. 1«. 21, 22) 



integument completely covered with dense spiculae. liototype: <y , Black Kill Conservation Park, South 



NEW CECIDOMYIIDAE FROM OLEARIA 



63 




Fig. 1. Sixth flagellomere of male Trigonomyia ananas sp. nov. Fig. 2. Head of male Trigonomyia ananas sp. nov. in frontal 
view. Fig. 3. Gonostylus of male Trigonomyia tulipa sp. nov. in dorsal view. Fig. 4. Gonostylus of male Trigonomyia 
cristata sp. nov. in dorsal view. Fig. 5. Wing of male Trigonomyia ananas sp. nov. Fig. 6. Genitalia of male Trigonomyia 
ananas sp. nov. in dorsal view. Fig. 7. Last three flagellomeres of male Trigonomyia ananas sp. nov. Scale bars = 100 
pro 1-4, 6, 7; I mm 5. 



64 



P. KOLESIK 





8 



10 





14 



^ 




15 




>r 



X»'* l **wWY*w*>t*f\ 



( 



( 


jj 






- _.. .__ 


-"«*&* . 


'{-■ 


l hi 


9 9 ' > 



16 



17 



Fig. 8. End of ovipositor of female Trigonomyia ananas sp. nov. in ventral view. Fig. 9. End of ovipositor of female Trigonomyia 
cristata sp. nov. in ventral view. Fig. 10. End of ovipositor of female Trigonomyia tulipa sp. nov. in ventral view. Fig. 

11. End of last tarsomere with claw, empodium and pulvillus of female Trigonomyia ananas sp. nov. in lateral view. Fig. 

12. Sixth sternite of female Trigonomyia ananas sp. nov. Fig. 13. Sixth flagellomere of female Trigonomyia ananas sp. 
nov. Fig. 14. Fifth tergite of female Trigonomyia ananas sp. nov. Fig. 15. Last three flagellomeres of female Trigonomyia 
ananas sp. nov. Fig. 16. Last two abdominal segments of larva of Trigonomyia ananas sp. nov. in dorsal view. Fig. 17. 
Head and first two thoracic segments of larva of Trigonomyia ananas sp. nov. in ventral view. Scale bars = 100 /tm 8-10, 

13. 15-17; 50 pm II; 500 ^m 12, 14. 



NEW CEC1DOMYIIDAE FROM OLEARJA 



65 




22 



23 



24 



Fig. 18. Trigonomyia tulipa sp. nov. - flower gall on Olearia axillaris. Fig. 19. Frons of pupa of Trigonomyia ananas sp. 
nov. Fig. 20. Trigonomyia cristata sp. nov. - flower gall on Olearia axillaris. Fig. 21. Trigonomyia ananas sp. nov. - flower 
gall on Olearia ramulosa. Fig. 22. Prothoracic spiracle of pupa of Trigonomyia ananas sp. nov. Fig. 23. Prothoracic spiracle 
of pupa of Trigonomyia cristata sp. nov. Fig. 24. Prothoracic spiracle of pupa of Trigonomyia tulipa sp. nov. Scale bars 
= 10 mm 18,' 20, 21; 500 ,mi 19; 50 pm 22-24. 



66 



P KOLESIK 



Australia (34°54'S, 138°44'E], 20.ix.1994, P. Koiesik, 
reared from flower gall of Oleario ramulosa (Labili. ) 
Benth., sampled I7.ix.1994, 121294 [SAM|. 

Allotype 9, same data, 121295 [SAM]. 

Paratypes (all sampled with holotype): \<J [SAM], 
I a IANJCI, 2 9 9 [SAM], 2 9 9 [ANIC], 2 pupal 
skins [SAM], 2 pupal skins [ANIC], emerged 
20.ix-8.x.l994; 2 larvae [SAM] t I larva [ANIC]. 

Other material (all sampled with holotype): 
3 9 91USNM], 4 pupae [SAM], 3 pupae [USNM], 
crneTged 5-8.X.1994; 4 larvae [SAM], 2 larvae 

(USNM). 

Description 

Male (Figs 1-2, 5-7) 

Colour: antennae grey, head black, thorax brown, 
abdomen with scleroti/.ed parts black and non- 
scleTOiLzed red (same in other two species) Wing 
length 3.1 mm (2.9-3.2), width 1. 1 mm 0. 1-1.2). 
Antenna totaJ length 1.5 mm (1.5-1.6)- Gonostylus 124 
fitu (12M27) long, 45 /an (4349) wide, length of apical 
claw of gonostylus 17 ^an (16-18). 

female (Figs 8. 11-15) 

Colour as in male Wing Jength 3.0 mm (2.9*3.1), 
width 1-1 nun (1.0-1-1). Antenna total length 1.4 mm 
(I.3-J.5). Cercus 65 fxm (60-68) long, 57 tan (57-58) 
wide, setae 5-28 pal long. 

Larva (Figs 16, 17) 

Colour red (same in other two species). Total length 
3.0 mm (2.7-3.6). Head capsule: length 57 tan (50-62). 
width 95 jim (92-101), posterolateral apodemes 13 fim 
llt-16) long; antenna 17 fan long, 6 fttti wide at base. 
Length of setae; 2-3 ;mi in ventral papillae \>l thorax 
and abdomen 1 VII. 10-20 /*m in remaining papillae. 
Intcgumcntal spiculae 1-2 pm long. 

Pupa (Figs 19. 22) 

Colour: non-sclerotized parts of abdomen red. 
remaining pans dark-brown (same in other two 
species). Total length 3.3 mm (2.3-3.9). Length of setae 
on cephalic papillae 361 am (354-369). Prothoracic 
spiracle 98 f*m (93-103) long, trachea 70 pm (60-75) 
long 

Oall (Fig. 21) 

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

Etymology 

The word "ananas", a noun in apposition, is the 
generic name of pineapple and refers to the 
resemblance of the gall to a pineapple. 



Trigonomxia cristata sp. nov. 

(FIGS 4. 9, 20, 23) 

Holotype: Cf a Beachport, South Australia [37°29'S, 
l40 J 00'E], 8.x. 1994, P. Koiesik, reared from flower 
gall of Olearia axillaris (DC.) F Muell. ex Benth-. 
sampled 6.x. 1994, 121296 ISAM] 

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

Paratypes (all sampled with holotype): 1 cf |SAM|, 
IC [ANIC|, 2 9 9 ISAM]. 29 9 [AN[CJ. 3 pupal 
skins ISAM), 2 pupal skins |AN(C|, emerged 
849.x. 1994; I larva [SAM]. 

Other material (all sampled with holotype). 2 9 5 
[USNMJ, 3 pupae [USNM], emerged S-19.X.1994; I 
larva [SAM]. 3 larvae [USNM) 

Description 

Male (Fig. 4) 

Wing length 34 mm (3.4-3.5), width I 3 mm 
(1.3-1.4). Antenna una! length 1.7 mm (17-1.8). 
Gonostylus 150 /im (145-157) long, 55 t an (50 58) wide, 
length of apical claw of gonostylus 20 pm (19-22). 

Female (Fig. 9) 

Wing length V2 mm (31-3.4). width 1.2 mm 
(1.1-1.2). Antenna total length 1.4 mm (1.3-1.6). Cercus 
60 urn (55-63) long. 50 a,m (47-52) wide, setae 17-45 
fim long. 

Lan-a 

Total length 3 mm (2.4-3.4). Other measurements 
within the range of 1\ ananas. 

Pupa (Fig. 23) 

Total length 3.7 mm (3.1-4.2). Length of setae on 
cephalic papillae 386 /im (361 427). Prothoracic 
spiracle 96 /im (86-107) long, trachea 57 ,un (51-621 
long. 
Gall (Fig. 20) 

Flower bud transformed into monothalamous, thick 
walled gall, 4-8 mm long. 3-6 mm wide, covered 
entirely with numerous, dcnsely-haircd, malformed 
ligulcs growing from proximal end When fresh, both 
ligules and gall wall green in colour. One larva in each 
gall Pupation takes place within gall. At Beachpon, 
T cristata was found with T. iulipa on the same shrubs 
with up to 20 galls of each species per plant. 

Etymology 

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

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

Holotype: cr, Beachport. South Australia [37°29'S, 
14XT00 El, 9.X.1994. P. Koiesik, reared from flower 
gall at Olearia axillaris (DC.) F. Muell ex Benth. ( 
sampled 6.x. 1994, 121298 [SAM]. 



NEW CECIDOMYDDAE FROM OLEARIA 



U! 



Allotype: 9, same data, 121299 [SAM]. 
Paratopes (all sampled with holotype): 2c c* (SAM|. 
lo- |AN]C], 2 9 9 [SAM], 2 9 9 (ANIC), 3 pupal 
skins [SAM], 2 pupal skins IAN1C], emerged 
9-17.x.l994 ? 2 pupae [SAM], 2 pupae | ANIC]; 3 larvae 
ISAM], 2 larvae |ANIC|. 

Other material (all sampled with holotype); I 9 
[USNM1, 4 pupal skins |SAM], 3 pupal skins 
fUSNM], I pupa |SAM], 3 pupae [USNM|. emerged 
917.x. 1994. 

Description 

Male (Fig. 3) 

Wing length 3.3 mm (3.3-3.4), width L3 mm 
(1.2-1.3) Antenna total length 1.7 mm (1.6-1.8). 
Gonostylus 121 /<m (119-126) long, 50 /mi (45-53) wide, 
length of apical claw of gonostylus 13 ^m (12-14). 

Female (Fig. Ill) 

Wing length 2.7 mm (2.4-3.2), width 1.0 mm 
(0.7-1. 1). Antenna total length 1.3 mm (1.1-1.5). Cercus 
56 ftm (52-61) long, 50 /on (43-55) wide, setae 6-20 
nm long. 

Larva 

Total length of the only specimen 2.7 mm. Other 
measurements within the range of ?! ananas. 

Pupa (Fig. 24) 

Total length 3.2 mm (2.8-3.4). Length of setae on 
cephalic papillae 395 M m (364-455). Prothoracic 
spiracle 51 fOQ (48-53) long, trachea same length. 

Gall (Fig. 18) 

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



Etymology 

The word tk tulipa T ', a noun in apposition, is the 
generic name of tulip and refers to the resemblance 
of the gal! to a tulip. 



Key to species of Trigonomyia 

! Trachea reaching end of thoracic spiracle in pupa (Fig. 
24). Apical claw of gonostylus diminutive, 1/4 Of 
gonostylus width (Fig. 3) From untuned galls oi Olearia 

axillaris (Fig. 18) - - T tulipa 

Trachea never reaching end of thoracic spiracle in pupa 
(Figs 22, 23). Apical claw ol gnnoMylus large, more than 
F3 ol gonostylus width (Figs 4. t>). From tufted galls ut 
Olearia spp. (Figs 20, 2J) 2 

2. Longest setae on female cercus shorter than 2/3 of cercus 
width (Fig. 8). From pineapple-shaped, glabrous, thin 
walled galls of Olearia ramutosa (Fig. 21). .T. ananas 
Longest setae on female cercus longer man 2/3 of cercu* 
width (Fig- 9). From hairy, thick walled galls of Olearia 
axillaris (Fig. 20)- - T. cristaui 



Acknowledgments 

The Ministry of Environment and Planning South 
Australia kindly gave permission to collect in the Black 
Hill Conservation Park. Abid A Munir Stale 
Herbarium of South Australia Adelaide courteously 
identified the host plant species. 1 am grateful to John 
D. Gray Department of Horticulture, Viticulture and 
Ocnology University of Adelaide and Raymond J. 
Gagne Systematic Entomology Laboratory USDA 
Washington DC USA for their careful review of the 
manuscript. 



References 



Cooke, D. A. (1986) Family Compositae tAsltsraeeue) pp. 
1423-1486 In Jessop, J. P. and Toelken. H- R- (Fds) Flora 
of South Australia, Part III <JNMcimniaceae-ComposiUu.'r 
(South Australian Government Printing Division, 
Adelaide). 

G.vJNk. R J. 1 1981 '» Family Cccidomyndae pp. 257-292 In 
MeAlpine, J. F . Peterson, B. V-. Shewetl, G fit. Teskey, 
H. J , Voekerofh, J. R & Wood, D. M. (Eds) "Manual 
of Nearcttc Dipfera I" (Canadian Government Publishing 
Centre. Quebec). 



(1994) "The Gall Midges of the Neotropical Region" 

(Cornell University Press. Ithaca. New York). 

Sobhian. R. & Ismono N, lift press) A review (il 



the genus Psfttrtwtm (Diptera: Ceeidomyiidae), Old 
World pesta of tiimuru, and description of Ihrcc new 
species. Isratt J. Ent, 
Koi r.siK. P. (1905) Asfthonriyiia datUmaea? a new species 
of Ceeidomyiidae (Dipteral damaging leaves and branches- 
of hop bush, Dodonaea \4xtoia (Sapindaceae) in Australia, 
Thins. R. Soe £ Ausi !L9, 171 176. 



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



Brief Communication 



Summary 

The Australian hylid frog Litoria raniformis (Keferstein) is a member of a group of 
similar species known as the L. aurea complex 1 , and is one of the largest species in 
Australia 2 (snout to vent length up to 104 mm). The geographic range of the species 
extends from South Australia through Victoria, the ACT and Tasmania to eastern 
New South Wales 3 . The species also has been introduced into New Zealand and has 
become established there 4 . 




Fig. 1 Pelvis of extant Utorta ranijormis from left, lateral 
aspect Length of ilium = 31 mm. (Photo: P. Kempster) 



Transactions of the Ruyat Society ofS. Aust. (1996). 120(2). 69. 

BRIEF COMMUNICATION 

FIRST FOSSIL RECORD OF THE HYLID FROG UTORIA RAN1FORMIS (KEFERSTEIN) 

The Australian hylid frog Utoria ranijormis (Keferstein) 
is a member of a group of similar species known as the L. 
aurca complex 1 , and is one of the largest species in 
Australia" (snout to vent length up to 104 mm). The 
geographic range of the species extends from South Australia 
through Victoria, the ACT and Tasmania lo eastern New South 
Wales 1 The species also has been introduced into New 
Zealand and has become established there 4 , 

It has been a source of surprise that such a large species 
seemingly is absent from Moloeene and Pleistocene sites in 
south-eastern Australia where other extant sympatnc species 
have been found in abundance". 

Here we report the first specimens of L ranijormis from 
the fossil record. The ilial descriptive terminology follows 
Tyler 6 . 

The largest and most complete specimen is a left ilium. 
located in March 1995. from material extracted at the East 
Low' site at Henschke's Cave. SA (Lai. 36 L "iR'-f)6, Long. 
140^45' ^8). The specimen has been deposited in the 
palaeontological collection at the South Australian Museum 
and registered as SAM P35305. The specimen has a length 
of 28.0 nun which is less than the known maximum ilial length 
of the species (35 mm) 5 . However, it is larger than other 
Pleistocene frog ilia known from the area and its identification 
has been confirmed by comparison with extant specimens, 
an example of which is shown in Fig 1. SAM P353U5 is 
fragde and partly encrusted with matrix, hence Ihe extant 
specimen in more useful for identification purposes. A line 
drawing of the sectional form of the ilium of this species has 
been published elsewhere 7 , 



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

We refer two other partial ilia from Henschke's Cave to this 
species: SAM P32249 and P35306. 

The age of the deposit has been estimated to be from 
35.00*/ lo 100.000 y years. These papers, piovide informa- 
tion on the deposit ional nature and siratigraphic sequence ol 
the material. 

We thank the Australian Research Council for funding 
Michael Tyler's investigations of fossil frogs and Peter 
Dempster for Figure I. 



'Tvler, M. J. & Davies, M. (1978) Aust J. Zool Suppl, 63, 
147 

: Tyler, M. J. (1978) "Amphibians of South Australia" 
(Handbooks Committee, Adelaide). 
Tyler, M. J. (1994) 'Australian frogs. A natural history" 
(Reed, Melbourne). 



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

-Tvler, M. J. (1977) Trans. R. Soc. S. Aust. 101(3), 85-89 

°Tvlei; M. J. (1976) Ibid 101(1) 314. 

7 Tvler, M. J. (1986) Alcheriiua 10. 40M02. 

*PJedge, N. S. (1981) Ibid. 105(1), 41-47. 

u Barrie, tt J. (1990) Mem. Qlii Mus. 28(1), 139-151. 



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



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



Brief Communication 



Summary 

Litoria revelata Ingram, Corben & Hosmer, 1982 is a medium sized tree-frog that has 
three disjunct populations; in northern Queensland (Atherton Tableland and the 
Bellenden-Ker Range), mid-eastern Queensland (Eungella Plateau) and the extreme 
corner of south eastern Queensland and northern NSW, Australia 1 . Herein we present 
a description of the tadpole of L. revelata from the rainforest in the Eungella region in 
mid-eastern Queensland. Habitat and life history notes are presented to assist 
identification in the field but these are intended as a guide only and tadpoles could be 
found in different habitats and months from those given. 



Transactions of the Royal Society of S. Ausl. (1996). 120(2), 71-73. 
BRIEF COMMUNICATION 



THE TADPOLE OF LITORIA REVELATA INGRAM. CORBEN AND HOSMER, 1982 

(ANUKA: HYLIDAE). 



Litona revektla Ingram, Corbcn & Hosmer. 1982 in a 
medium sized tree-frog that has three disjunct populations; 
in northern Queensland (Atherton Tableland and the 
Bcllenden-Ker Range), mid-eaatern Queensland (Eungclla 
Plateau) and the extreme comer of south eastern Queensland 
and northern NSW, Australia 1 . Herein we present a descrip- 
tion of the tadpole of L. revelata from the rainforest in the 
Eungella region in mid-eastern Queensland. Habitat and life 
history noies are presented 10 assist identification in the field 
but these are intended as a guide only and tadpoles could be 
found in different habitats and months from those given, 

Tadpoles were collected in November and December of 
1993 at several stream sites near the Eungella township. 
approximately 70 km west of Maekay, central Queensland, 
Australia (Table I) A sample of larvae was preserved in 10% 
formalin and others were reared to metamorphosis tor 
identification, Terminology follows Altig* and Hero ; 
developmental stages follow C»o*ner . Measurements were 
taken using vernier callipers. Height ot the caudal muscles 
and tins was measured at mid- Length ot the tail. The drawings 




Fig, I. Tadpole of Lloria rcxrlata (QM J 59240; Gosner slaye 
35; TL 31.5 mm). Scale bar = 9 mm. 



depict melanin patterns that persist in preserved specimens 
(10% formalin). The colour descriptions should be treated 
with caution as tadpole colour is often a function of water 
clarity 5 . Drawings were made of two representative 
specimens (Figs 1 and 2) placed in the Queensland Museum 
Brisbane (QM J 59239 and J 59240). The labial tooth-row 
formula (LTRFj is based on observations of all specimens 
collected at Gosner 4 stages 25 through 45 (QM J 59241 and 
J 59242; Table I). 

Desiiiptitm: Eyes lateral; eye diameter 14.5% of the body 
length for stage 35 tadpoles and 14.7% for stage 40 tadpoles 
Nares dorsal, nearer u> tip of snout than to anterior edge of 
eye; nanal margin without rim. spiracle paragyrinid (Fig. 2 
C: located well below the horizontal longitudinal axis but not 
un the midline so neither sinistral nor medioventraJ is entirely 
applicable 6 ), unpigmented, opening directed posteriorly. 

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

In lite, body opaque, appearing "bluish" and heavily 
pigmented with lighter pigmentation around eyes, darkly 
pigmented supracranial patch (especially in larger tadpoles i 
extending posteriorly over spinal cord (Be. 2H); distinct broad 



'abm: I. Dotes* btCttitUe\ and Museum Numbers for additional specimens examined in this study: Ml William (upper Cattle 
V.; Map S65X MGR 666740). Mi Puvid (upper Cattle Cn; Map 8655. MGR 678744K 



Tabit. I 
C 



Date 



Place 
Collected 



Gosner Stage 


Body Length 


Tbtal Length 


(No i 


(mean) 


(mean) 


25 


4.6-8.1 


11.2-38 J 


(7) 


(6.6) 


[13.31 


2(. 


9.5 


23.2 


27 


10.6 


26.7 


28 


8.4 


21.5 


29 


11.3 


27.4 


n 


12.0 


29.8 


33 


11.7 


30.6 


^o 


12.4 


312 


37 


13.3 


33.3 


42 


11. 1-12.0 


29.6-353 


(2) 


(11.5) 


02 J) 


43 


10-5 


- 


44 


U .4-13.4 


- 


15) 


(12 3) 




25 


8.8-10.6 


21 1-25.2 


(3) 


(97) 


(23.1) 


27 


II 4-11 *> 


26.1-27.6 


S 


(ll.b) 


(26.8) 


12.0 


33.0 


37 


14.6 


431 


41 


14.4 


40.8 



Qd Mus. No. 



27.x i.93 



Cattle Cr, 
Mt William 



J 59242 



2V.xi.93 



Cattle Cr. 
Ml David 



J 59241 



72 









B 











C 










D 




df 



Fij: 2. Tadpole of t/fcwwi rcveluta (QM J 59239; Gosncr Stage 40; TL 40.8 mm), A. Oral disc. B. Dorsal view. C. Ventral 
view. D. Dorso-lateral view. Scale bars = I mm (A.), 5 mm (B,C,Dj. 



73 



vertical subdermat lines on dorsal side of each nari*- 
Horizontal bund or patch from snout lo eye (Fig. 20). 
Pigmentation often lighter during earlier stages (Fig. I) than 
at later stages (Fig. 2B-D). In ventral view intestinal mass 
visible, intestinal coils partially visible and obscured by heavy 
pigmentation; branchial region semi transparent. Tail 
musculature an even shade of grey/brown with additional 
mclanophorcs concentrated dorsally (Fig. 2D). Dorsad and 
ventral fins transparent, with even stippling of dark 
melanophorcs. often outlining venation. 

A tadpole at Stage 35 »Fig. U bad the following 
measuienicui* (inmi: total length 31,5. body length 11.7, body 
width 6 5, bixJy height 6.0, tail height 7.-. interorbiial distance 
.S.t), internarial distance: 2.1, eye -nans distance 2.0. A tadpole 
at Stage 40 {Fig 2) had the following measurements (niuo: 
total length 40.8, body length 14.3. body width 8.2, h»dy height 
7.3. tail height 8.3. inierorbital distance o.2, internarial distance 
2.0, eye-nans distance 2.6. Tadpoles vary in total length from 
II 2 mm at Stage 25 to 43.1 mm at Stage 37 (Table l|. 

Diagnosis At the sties studied, live tadpoles of/., rewittm 
can easily be confused with L chtoris as both species occur 
in mid-water sections ot isolated streamside pools and they 
have similar body shape and oral disc formula. Live tadpoles 
of/., rexehta have a bluish sheen covering (he intestinal mass 
tod the intestinal coil is partially visible (Fig. 2C). In contrast, 
/.. clitoris has a golden sheen covering the intestinal mass, 
the intestinal coils arc visible mid -ventral ly and golden 
ehromatophores cover the heart. 

Ill preservation, tadpoles o\' L revvltiut have pigmentation 
coveting the intestinal mass making the intestinal coils only 
partially visible. In contrast, L. thloris has a transparent 
veniral surface and the intestinal coils are clearly visible The 
position of the spiracle, paragyrimd in L mr/wwand sinistral 
in L fJiIori.y and the dark pigmentation on the oral papillae 
ol /- revt'faUi (with only few scattered pigments on the oral 
papillae of £, chloris) also distinguish these two species. 
Interestingly, we only know of one other Utoria sp. in 
Ausualia wirh a paragynnid spiracle it. rahetla, unpubl). 
Tadpoles of/,. rewUmx were found in sympairy with tadpoles 
ot L thtitri.s mul TauJactylus liemi. Adult frogs of T. 
mngplhnsh and MLxophyex fasciolatus were also observed 
in adjacent streams 



Jiabitui: Tadpoles of L. revvlata were found in isolated 
bedrock pools adjacent to fast flowing rocky streams 
surrounded by pristine rainforest. Each pool contained leaf 
litter and algae and was between 1.5 and 2.5 m from the 
stream. No fish were observed ot captured by dip netting the 
[tools. Water temperatures were noticeably higher in the pools 
than in the adjacent stream (Table 2) Pool dimensions in 
November were 100 em x 50 em x 10 cm deep for pool I 



Tabu: 2. Wbttr temper at tin's C C) oj pooh atul the itdjtueni 
\trcfiin /Mv» vrnv. 



Site 


Dale 


Vonl 1 


Fool 2 
10.0 


Adjacent Stream 


Cattle Cr 


JB.ix.94 


i4.9 


142 


Ml William 


16. \ 94 


I6.K 


17.2 


IS-2 




13. x» .04 


19.7 


20.5 


17.9 




ll.xii 94 


20.0 


106 


18.0 


Cattle Cr 


JU.ix 94 


175 


160 


13 5 


Ml David 


17*04 


21.3 


- 


15.0 




I4.\i.y4 


20,0 




I7JJ 




12 xi. .94 


2:. 5 


22.8 


i7.9 



and 200 Cm x 100 em x 25 uo deep for pool 2. Tadpoles 
were generally observed in the midwaier and surface water 
rather than the benthic layer of the water column and were 
frequently observed rising to t»»e surface to gulp aii 

This research was partially funded by the Australian Nature 
Conservation Agency, the Queensland Department of 
Environment and Heritage, the Wei Tropics Management 
Authority and the Cooperative Research Centre for Tropical 
Rainforest Ecology and Management. Research was carried 
out under a Qld Dept ol Environment and Heritage "Permit 
to Take' 1 no. '100177. Wc wish to thank Michael Cunningham 
lor his contribution to (his paper and Julie Martin who 
prepared the illustrations and the volunteers who assisted in 
the J.C.U, Eungella Frog Search in Novcmhcr/December 
I99A Ross Alford provided logistical support at J.C.U. and 
Marion Anslis gave valuable comments on a draft. 



'Ingram, G- J. t Corhcn C. X & Ilosmer, W. 1 1982) Mem. 

Qd Mus. 20, 635-037, 

r AJtfgi R. (1070) Hcrpctologica 2ft, 180-207. 

^Hero, .l.-M. (IOOOi Arna/oniana U, 201 262 



4 Gosner, K. I, (I960) Herpeiologica 16. 183-190. 

-"Bragg, 4. N. (10571 Copeia 1957. 36-39. 

Mohnston, V. h, & Altii?, R. <I9K6) Herp. Rev. 17. 3b -37. 



lEAK MA.RC HERO Wet Tropics Management Authority CRC Tropical Rainforest Ecology and Management Dept of Zoology 
James Cook University Townsvillc. Qld 4811. SHF.RHt: FICKLING & RICHARD Rl.TALl.JCK Dept of Zoology James 
Cook UniversitV Townsvillc. Qld 4811 



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

14.vL1907-9.v.l995. 
President of the Royal Society of South Australia Inc. 1961 

Summary 

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

I first came across the name N. H. Ludbrook when I was a student at the University of 
Adelaide in the late 1960s. While I was searching through the stacks in the Barr Smith 
Library on some aspect of the evolution of interior deserts, her name appeared a 
number of times in a paper dealing with geomorphology. The more I searched related 
papers the more her name recurred. I must confess that, then, I didn't know 7 whether 
N. H. Ludbrook was male or female. All I knew was that the name was referred to in 
an array of papers dealing with stratigraphy, geological evolution, palaeontology, 
palaeoclimate, ancient glaciations and the list went on. And it didn't seem to matter 
w ; hat part of the Phanerozoic either. I admit I thought that a person touching so many 
aspects of geoscience had to be of great physical and scientific stature. It was not until 
twenty odd years later when I actually met her that I found I was wrong on one count 
but I was certainly not disappointed. What a marvellous person and an extra-ordinary 
scientist I found her to be. 



74 




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

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



Transactions of (he Royal Society of 5*. AusL (1996). 120(2). 74-77. 

OBITUARY 

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

14 vi. 1907 - 9.V.1995. 

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



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

I first came across the name N. H. Ludbrook when 
1 was tt student at the University of Adelaide in the 
laic 1960s. While I was searching through (lie slacks 
in the Ban Smith Library on some aspect of the 
evolution ol interior deserts, her name appeared a 
number of limes in a paper dealing with 
gcomorphology. The more I searched related papers 
the more her name recurred. 1 musl confess that, ihen, 
I didn't know whether N, H. Ludbrook was male or 
female. All I knew was that the name was referred 10 
in an array of papers dealing with stratigraphy, 
geological evolution, palaeontology, palaeoclimate. 
ancient glaciations and the list wcnl on. And it didn't 
seem to matter what part of the Phanerozoic cither. 
I Admit I thought that a person touching so many 
aspects of geoscience had to be ol great physical and 
scientific stature. It was not until twenty odd vcars later 
when ] actually met her that I tbund 1 was wrong on 
one count but I was certainly not disappointed What 
a marvellous person and an extra-ordinary scientist I 
found her to be. 

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

hollowing her marriage in 1935, she and her 
husband, Wallis Verco Ludbrook, moved to Canberra 
where, undaunted, she continued her interest in Camo- 



zoie Mollusca. It was fortunate at this lime that the 
Commonwealth Palaeontologist function was moved 
from Melbourne to Canberra, undoubtedly facilitating 
her continuing interest in palaeontology. While in 
Canberra, from 1942 to 1949, Nell worked as Assistanl 
Geologist in the Commonwealth Bureau of Mineral 
Resources dealing wilh statistics of strategic minerals. 
In 1950 she travelled to London. Here at Imperial 
College and as a visiting scientist at the British 
Museum (Natural History) she continued CD extend her 
palacoiitological studies. Nell was awarded her PhD 
in geology (1952) from the University of London and 
the DIC in palaeontology for research on Pliocene 
Mollusca from the St Vincent Basin Out of this 
research developed an aulhorative chapter on fossil 
>ciphopods in the first edition ot the Treatise ot 
Invertebrate Paleontology"* (I960). 



<*?**rr 




*!■ 



Sampling Barly Cretaceous Eromanga basin sediments near 
Marree. 1963, Photograph by B. (J Forbes. MESA photo 
no. 20035. 



76 



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

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



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

"Retirement" really meant the continuation of her 
love of geology and especially for fossil Mollusca. She 
worked as a consultant in palaeontology to the 
Department o\' Mines and Energy unlil 1993, at which 



t 




N. H. Ludbrook and J. Spcncc examining Cainozoic sediments at North West Bend, along the River Murray Photograph 
by A. R Crawford. MESA photo no. T002001. 



77 



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

Although the vast number of her publications (over 
70 scientific papers and monographs) and Government 
reports was known to me, I only became aware of the 
full extent of her extraordinary energies whilst i was 
researching material for the J995 Volume 2 of the 
"Geology of South Australia" During the course of 
rummaging through filing cabinets in the 
Biostratigraphy Branch containing countless numbers 
of her Report Books 1 came across a huge number ol 
unpublished letters and personal communications to 
geologists in companies, academia and government 
carefully outlining the results of work undertaken for 
them, each almost of quality to be published notes in 
their own right We arc now the custodians of Nells 
journals, books and notebooks, donated by her in 1994 
and now housed in (he N. H. Ludbrook Memorial 
Library at Mines and Energy South Australia. 

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



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

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

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

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



NEVILLE F. ALLEY 



OBITUARY 
STANLEY JOE EDMONDS, BA, BSc, MSc, PhD, Dip Ed. 

13.ii.1909-16.vii.1995. 
President of the Royal Society of South Australia Inc. 1965 

Summary 

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

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



78 





STANLEY JOE EDMONDS 

BA, BSc, MSc, PhD, Dip Ed. 

Photograph courtesy of the SA Museum 



humutiitms of the &>yot AV'cv/'/v of S, Ami. (1996). 120(2). 78-82. 

OBITUARY 

STANLEY JOE EDMONDS, BA, BSc, MSc. PhD, Dip Ed 

13. ii. 1909 - I6.vii.l995. 
President of (he Royal Society of South Australia Inc. I96S 



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

His working life fell roughly into three periods each 
of about iweniy years" duration. He was a SCftooi 
teacher at Adelaide High School from 1931 to 1952. 
he (hen taught and conducted research in the Zoology 
Department of the University n\' Adelaide from 1952 
to 1^74 and finally, as an Honorary Associate at the 
South Australian Museum, he continued his research 
from 1972 to 1995 

Stanley Joe Edmonds was born in Adelaide (Soudi 
Australia) on |3 February, life?. He attended the 
Thebarion Primary School from 1915-1922 and the 
Woodville District High School from 1923-1925, 
Obtaining his Intermediate Certificate in 1924 and his 
Leaving Certificate the following year Jn 1926 he 
joined the Lands and Survey Department of the South 
Australian Public Service with the intention of 
becoming a surveyor. During this time he Studied 
science part tune as a private student at the University 
of /VJeiaide doing Mathematics i during the day and 
Chemistry I and Physics 1 al night 

lit 1927 he joined ihc Education Department and 
entered the Adelaide Teachers College. He graduated 
in 1929 with a BSc majoring in Inorganic and Organic 
Chemistry arid began his leaching career in 1930 
teaching for six months at Woodville High School. 

In 1931 he began his impressive twenty-year 
association with Adelaide High School, ultimately 
becoming a Special Senior Master in Chemistry and 
Oencral Science and teaching Leaving Honours 
Chemistry from 1945 to 1951. During this time, he 
obtained three further degrees - a BA in 1935 in which 
he majored in Latin and English, a First Class Honours 
in /oology in 1945 (after completing Zoology L P and 
III in I94L 1943 and 1944 respectively) and an MSc 
in 1947 

However, it was his broad interests and sense of 
humour combined with his great teaching ability that 
endeared him to his students. He was interested in 
sport, particularly tennis and hockey at which he 
excelled, acting, singing and the arts 



During his last few years at Adelaide High School 
Stan became interested in Zoological research and 
began a series of collaborative studies with T Harvey 
Johnston, the Foundation Professor of Zoology at 
Adelaide University and a noted Parasitologist These 
studies on Australian Acantboeephala (spiny -headed 
worms parasitic In the alimentary canals of various 
fish, birds and mammals) were first published in W : 
and continued for several years after Professor 
Johnston's death tn 1951. During this period Stan 
widened his interests to include free living niarnu 
organisms and in 1948 had a paper published on "Un- 
common species ot animals and their distribution on 
an intertidal platform at Pennington Bay. Kangaroo 
Island'*. These mounting interests in Zoology led in 
1952 to his resignation from the Education Department 
to take up an appointment as a lecturer in the Zoology 
Department. University ol Adelaide under the newly 
appointed Professor of Zoology, W P. Rogers, an 
authority on the physiology and biochemistry ul 
parasitic nematodes. 

It was at this time that 1 had the pleasure ol meeting 
Stan as I had been appointed to the Zoology 
Department as a Demonstrator earlier that year. One 
always associated him with laughter or at least a smile. 
His miming of sewing his fingers together and then 
threading the needle through various parts of his ami 
so thai (be whole could be moved mechanically by 
pulling on the thread, was always demanded of" him 
al departmental parties and was always accompanied 
by gasps of horror from the faint-hearted and much 
amusement from the initiated. 

Later, in his chapter on Zoology in "Ideas and 
Endeavours. The Natural Sciences in South Australia" 
Stan desej ibed what a busy time his early years in the 
Zoology Department had been as student numbers 
were increasing rapidly and Rogers was building the 
Zoology Department- Needless to say. in this chapter. 
Stan gave himself scarcely any mention. 

In addition to broadening his taxonomie interests, 
describing new species of Australian marine 
invertebrates, notably sipunculans and echiurans, Stan 
began to conduct a range of physiological and 
biochemical experiments on them. These physiologicaJ 
experiments were extended to the parasitic aeantho 



XI) 



cephalans and to the species Moniliformis duhius in 
particular. This species wk maintained in il»e 
laboratory in cockroaches, the intermediate host, and 
rats* the primary host. The distinctive rustle and odour 
of Stan's experimental cockroaches, as one entered the 
constant temperature room in which they were boused, 
are sounds and smells not easily forgotten! The 
nutrition and egg laying of these animals were studied 
and reported upon and, together tvitfi B. R- Dixon, a 
paper was published in Nasure nn Ihe uptake of small 
particles Through the body wall of M dubius. 

Around this time Stan collaborated with H. B S 
Wuuicrstey in what was the first significant paper on 
the inrertidal ecology of South Australia. It w*> also 
the fust paper published on this topic tn Austral ia that 
dealt in equal detail with both flora and lamia. 
Piirthermore, it dealt with the relatively sheltered 
coastline of South Australia which, with its gulfs- and 
bays and Kangaroo Island, differed front the more 
exposed coastline of the Eastern States. In tltc h&ht of 
then work Womcrsley and Edmonds were able to 
supply evidence, previously unavailable, for die 
btugeographical nomenclature of the southern 
Australian coastline 

In addition to his researches on irttertidal ecology. 
Scan also worked with Marian Specht on ecological 
studies of heathtand mi the Keith region of South 
Australia Thrs work involved monthly visits over a 
period of three years between 1952 and 1954 and 
resulted in Ihe accumulation of a vast amount of 
information that permitted judgements to be made on 
the faunal rhythms of hcathland in South Australia 

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

Stan Edmonds' work 06 the Public Examinations 
Board, a member from I960 to 1974, Chief Examiner 
in Biology for ten yean and deputy Chairman from 
197!V 1974, was a measure of the regard in wtuch his 
teaching experience was heW. 

Stan was to continue to undertake research on the 
slpUDeulans and the somewhat similar echiunins fiw 
many more years. In 1972 he co-authored a book wwh 
his late friend Dr A. C. Stephen of the Edinburgh 
Museum emided "The Phyla Sipuncula and Echiunf 



At the time of Dr Stephens death, much remained to 
be dooe and it was recognized that Stan was the only 
person who had the scholarship and energy to complete 
thia task. His share of this important contribution to 
marine studies was a large one, bringing information 
up to date and checking descriptions, records and 
translations with original specimens and daia. He 
arranged species into genera, provided keys for 
identification and was alone responsible for the sixty 
lull page illustrations. Some 320 species of sipunculans 
and BO species of echiurans had been described at the 
time this 527 page book was published by the Trustees 
of the Bnush Museum (Natural History} London. It 
was the first systematic monograph of the two phyl.i 
to be published this century and is likely to remain 
the standard reference work for many years to come. 

Stan retired from the University of Adelaide in 1974, 
having been made a Reader in /oology in 1973 He 
became an Honorary Associate of the South Australian 
Museum and over the next twenty years published & 
lurther thirty papers including several chapters in 
books He was a strong supporter of the Royal Society 
<rf South Australia Inc.. occupying the positions of 
Council member, Secretary, Vice President and 
becominp President in 1965. In 1982 he was awarded 
the Society's Sir Joseph \ferco Medal (or Ins 
distinguished scientific researches. 

In conclusion I quote from C. M. Ward MA, a Latin 
tcacliet and scholar of high repute and, at ihe time. 
Ac-tiny, Principal of Adelaide High School who wrote 
on 17 September. 1948 the following WDfdfc. u Mr 
Edmond? has a lively, genial personality, a good seme 
of humour and a resourceful cultured mind. He h of 
strong* independent character but always friendly and 
unassuming His honesty and integrity are 
unquestioned H A most fitting tribute to a much liked 
and respected personality who maintained these traits 
throughout his life. 

Stan Edmonds is survived by his wife Barbara fnee 
Hy) and a daughter Elizabeth. 



MAN F. BtRD 



Bihltograph} 



1947 Australian Ac&nthocepruii No. 5. Trans, ft &c S. ,+& 
71, 13 19. (With T. H. Johnston). 

t047 Australian AcanuWephala No. 6. Bee. S. .Aust. Mux. 

8. 555-562 (With T. H. Johnston). 
IWl Australian Acamhocephala No. 7. Trans. H. So*\ & Aim. 

11, 69-76. (With T. H. Johnston). 

1948 The commoner species of animals and their distribution 
on an intenidat platform at Pennington Bay, Kangaroo Island 
South Australia. Ibid. 167-177. 

H*51 Australian Acanthoccphala No. 8. find 74, l-V (With 
T. H. Johnston). 



1952 Australian Acawhocephala Nu. 9. ttrid 75, 16-71 <Wllh 
T. J*i. Johnston). 

1952 Marine zonalion in Australia in relation to a gen«™i 
scheme of classification, /. Ecol, 40, 84-90. (With H. B. 
S. WomereIey», 

1953 Acanthoccphal3 from Auckland and Campbell Islands. 
fee. Pom. Mas. N.Z. 2, 55-61. (With T H. Johnston) 

655 Australian Sipunculoidea l. The genera Sipunculu.\. 
Xrn* •-..',"'!• m ; and Siphonosoma. Aust, I Mar Frrshw #m. 
*. 82-97 



(955 Acanthocephala collected b> the Australian National 
Antarctic Research Expedition on Heard Island and 
Macquarie Island during 1948-1950. Tranx. ft. Sof. £ Ans( 
78, 141 144 (With T. H. Johnston). 

1956 Australian Sipunculoidea 2. The genera rVwscolam wia, 
Dcndwsumuim, Golfing, Aspidosiphott and Cloeoaphiu. 
Auxi. J Mar. Frrsiuw fifes 7, 281-3)5. 

1956 Chlormities of coastal warcrs m South Australia. 'fonts 
ft Sat S. Au>t. 79, 152-16* (With I, M Thomas) 

1957 The respiratory metabolism ot &4nifctttytF?iHffl 
<v/w/«Mrw Edmonds, iSipuneuloidea). ,'tiW. J Mar. 
frt'.vMt: Res, S, 55 63 

N57 Iht catabohsni of ttipogen compounds in DMdrrtfffi#pii*\ 
rymtxla&ttt 1 Edmbryfc f$piinauwdett). /few m-iv> 

IW Aeanthocephalafl'l.A'.Z.Afti: /i/imr. Jta tawrf. 
W29-19J1 /&y. B6, 92-98 

l»>57 Australian Acanihoeephala Nix 10. Hjwu, ft. Su V 

>4UH 80. 76 80. 
958 A general account of the inienidal ecology of South 

Australian coas-1*. Us/, J. Mor Freshw. Rex. 9. 2P-260 

tWuh H B S. Womersley). 
WW) Some Australian echiuroids (Echiuroidea) Trans ft. 

Sett S. .-i«.r/. 85, 89 ys. 
WoO Sipuncutids. from New Zealand and Chatham Maud 

N.Z. jOffp; 5'cv. & Must. Rt$, Bull 139, f 59-167 
[961 OrtX^iim7///£yd^nt t t4.v&iirdlSipuncu1nnk3t> ,*0w. Mt*x, 

■'tat. km U, 217-220. 
1962 Some notes on die abundance, cm imnment and nutrition 

rt'SipuncuhtsftuiUis L. at Morgat. BriQanv CA Itioi nun 

3, town 

i l »f) ! I wo !R-w echiumids (Ecfuuruidea) from Australia fraaf 

A' Sbc f Ajtf, S7 243 247. 
1964 Australian Acamhocepnaia No. \\. tbut 85, *l 95 
W» -5ipuno.il' .idea • d the Rims Sea, N. /. ttatf. SW. ,y fcjfeij 

/Cp.t fiW/ 167, 27-34 
I9fi3 Si/me e-Apenrnenls on the nutrition of /tfjni/ftjrab 

JuhUr, Meyer j Ac.tmhocephal.il, fomsit&ingy 55. 337 31 J 
1966 StphonuApflm fumaienxe a new sipunculoui tunn Hmwjjii 

(SipnriLuloideai Aft* Sri. 20, 3K6-3KS. 
196<. Uptake of sttulJ particles by Moniliformis ttitbius 

tAcanlrtotephala). Njturt (Loudon) 21)9, $> {Widi B I? 

IhV.MIJ. 

1'Jf,Vi Hatching tA the eggs, uf Motnhfitrmi-, { juh\us Mcvt'r. 
»/>//. Purasuoi, 19, 216-226. 

1966 SipuoculiMdei and Eehiumidca. Port Phillip Survey 
19579. Mem Nm. Mux. Melb, 27, 175-178. 

1967 iUruciinthnrhyniiws gtdoxiaxiU a new genon jiij SpcCw^s 
of Aeimhoccphab from fish AuslrahaD Acanlhivepliala 
No 12. ftflKfc ft, 5or. 5. .4Al$t. V], 4M3 

lOftK DiMribuiion of' selected groups of marine invertebrates 

in waters south dT35*S lar. Tolto It Antarctic Map t^olio 

Scries, pp 23-24. 

Stpuncula and tichiura. Amcr. Gcngrah. Soc. 
I%V Measurements of the osinolic pressure in ihe rwbnai of 

B)l\mor)>futs minuUts ( Acanlhocephitai in ihe irtesiine of 

the domestic duck. J, Ktp, Biol, 50, 69 77. tWilh D. W. 

T. Crompton) 
197| Some sipuncularb and echiurans chiefly from Guam 

fSipuncula and Echiura), Mivmnesita 7, 137-151. 
197! Australian Acanthocephata No. 13- Three new species- 

trasa, ft. Snc s fan. 97, 10-21 

1972 Marine mvertebrarx-N from Adelic Land, collected by the 
(2ih and 15th French Anlarclic Expeditions. 5. Sipunmla 
Teifws. Supp. 4. 83-86 

1972 "The phyla SipuncuU 3rd Rchiura" (Trustees British 
Museum fNai. Hist.), London). (With A. C Stephen). 

1973 Australian AeanuWephala No. 14. On two species of 
BmirhadinorftmchuA, one new. Trans, ft. She S. Ani ^7 
19 21. 



HI 

1973 A new spxies and genus of earthworm iMegaseoleeidac 
Oligochaeta) fn)m South Australia, tbid.2^21. (WioS B 
G. M Jamieson). 

1974 A new species of Sipuncula \A.\pidnvipkan e.xiyuu.s} 
belonging to ihc interstilia! fauna of marine beaches, 
collected by Mr L. Botosanaenu during the second Cobo 
Romanian Biospeleologjcal Bnpedition to Cuba, 1973. hi: 
J. Speli-ot, 6. 187-192. 

I97ti Respiration in sipuneulaos. Proc. [nlvntational S>rtip. 

Biol Aipuncuia *n\S Echiura. Kotar (1970] 2, 3 9. 
1976 Three sipunculan species (rwe new) ttmn New Zealand 

N.Z, J- Mar- fftshm ftr-A. 10. 217 224. 
1979 tnlcrtidal invertebrates pp. 65-16? fa Tyler, M. J. 

TwicLUc C R. & l fog, )_ K, (Eds) "Natural HiMo,v ol 

Kangjro4> Inland" 1'R. Soc, h> Ausl, LftC , Adcljiulr) 1 With 

1 M, Thomas), 

1979 Intertidal ecologv ot marine orpliisrus pp_ 167177. !h>,i 
(With H. B S Womerslev 1 ' 

1980 A revision of the svtetrwtics of AuNiralian sipuncu!an> 
tec. 5. Ausi. Mti&. 1». W4- 

1981 Dark Iniand hcathland, Sitmh Australia: fuunai rhjfhrro 
pp. 15-27 in Spccht, R, L. (Ed.) *Rci>syMems ot Ihc W>rld. 
9R Heathlands and Related Shrublands. Analytical Studio.' 
(Elsevier. Amsterdam). <Wilh M. M Spechi). 

I9S2 Sipuncul-anspp 299-311 hi Shepherd, S. A & TftOm« 
1. M. fBdsQ "Marine invertebraies uf Southern Australia 
Part I Handbook of Flora nnd Fauna ol South Australia' 
tGovi Printer, Adclaidt). 

|U82 Ectimraos PP _ 3 12 318. ttui 

19H2 AuNiralian AcandiocephalaJNo. 15. Four vpeciev Tmnx 
ft. Soc, $. Aust. 106. 7» 76. 

1982 Eehiura pp. (»5-66 ht "Synopsis am I l- Ofil^aitOil t'f 
Livinp Orgdni.vm.'." (McGraw-Hill. New Vorkl 

I9N2 A sipunculan. rep*->ned to i>_ 'n.invoii;)!" hutn itii. 

Nahtrhtnds ApeiIIci. P(/sfrw«i Pfrnfcawrff 52. 22> 2 w > 
I9K4 Phylum Sipuntula and Phvlum V,< htllnl pp. M 62 In 

Mather P. & Btnnwi. t iEds> "A Coral Beef Handbook" 

(Great Barrier Roct Society Bru-tuner 
|9H5 A icw -.jii L ie> of tVUi>co{t,\<m»i (StplfnvJIjd ln "" 

Australu TrtM$ H Stu s fcpi itN. 43 44 
I9H5 A new ectimran ShiUtfirUl fafftmintt (hrliiuro: 

Bi Tieilndae 1 iroto Sr\>, Xtaland and a note on New ftsiUftd 

evhiurum. fV.^. 7 A/aA fnfttftW Ar>^ 19, M)I-604 
;9R6 Sipuncula p. 2(W fa BoU^aneanu. L, (Ed > 'Stagflfcainu 

Mum!» 96" ■ {Brill & Backhuw Fctdcn), 
l$8fi A checklist of hclmrrrthv fmiT Australia!' birds. At. 

S jdttSJ WWl 19. 21V -325. (With P M Mawwn ^ L M. 

Ang^O 
1986 A note on s^me siptincuttM ftnm the Wcrthem Territory 

AustraJia. Jhv Bcagfa 3, 7-9 

1986 Zwtogy pp f62-2!2 M Twtdafe, C R , Tyler. M J 
di Davies, M. (Eds> "Ideat- and Endtavours. I Ut Natural 
Sciences in South Australia' (K. Sot, S. Aum. fnc, 
/Vdelaioe). 

1987 The sipunculan fauna (SipuneuJaj of Western Australia. 
6Vc W. Aust Mm. 13, 215-224 

I9K7 A note on the occurrence pi tiolhusottui rapUatuw 
(Linstow. 1880) (AcanthoccphaJa) koto j toJsc kilier whatc 
stranded on (be coast of Western Australia, Ibid. 317-318. 

1983 Obituary: I M. Thuniax. Her. 5, %W Mus, 21. 6J-6^ 

1987 Echiurafie from Australia, fbkt. 119-130. 

W87 Sipunculans and Echiurans. pp. 185-212 In Devaney. D- 
M. & Eldndge, I G, (Eds) "Reef and Shore Fauna oi 
Hawaii. Section ¥ (Bishop Museum Press, Honolulu* 

1988 .Aiwnitifortms .wmoni (linstow; 1898) n. gen., n- CDtflb. 
(Acanthocephala: Moniliiorniidac) from in;irsupials hi 
Australia and New Guinea. / ftimsitol. 75, 215-217. (With 
G. D- Schmidt). 

J989 A lisl or Australian Acanthocephala and their hosts. Hvc 
5. Aim, Muy 23, 127-133. 



82 

1991 Sipunculans and echiurans from the Phillipines and New 1992 A note on Phascolosoma turnerae (Rice) (Sipuncula). 
Caledonia (Estase 2, Musorstom 3 & 4). Mem. Mus. Nat. Ibid. 151. 

Hist Nat (A) 151 83-90 1993 Sipuncula and Echiura pp. 94-96 In Mather, P. & 

Bennett, I. (Eds) "A Coral Reef Handbook" (Australian 

1992 A new species of Acanthocephala from the green back Coral Reef Society, Brisbane). 

flounder Rhombosolea tapirina Gunther. 1862. Trans. R. 1993 Index to the Transactions of the Royal Society of South 

Soc. S. Aust. 116, 35-40. (With L. R. Smales.). Australia, Vols 102-113 (1978-1989). (With L. W Parkin). 



VOL. 120, PARTS 3 &4 
29 NOVEMBER, 1996 




Transactions of the 

Royal Society of South 
Australia 

Incorporated 



Contents 

Anstis, M & Littlejohn, M. J. The breeding biology of Litoria subglandulosa 
and L. citropa (Anura: Hylidae), and a re-evaluation of their 
geographic distribution --------- 83 

Dyson, I. A. Stratigraphy of the Neoproterozoic Aruhna and Depot Springs 

subgroups, Adelaide geosyncline - - 101 

Dyson, I. A. Stratigraphy of the Neoproterozoic Tent Hill Formation and 

Simmens quartzite at South Tent Hill on the Stuart Shelf, 
South Australia - - - - - - - - - -117 

Taylor, G. S., Austin, A. D. & Davies, K. A. Biology of the eucalypt 
gall-forming fly, Fergusonina flavicomis Malloch (Diptera: 
Fergusoninidae) and its associated hymenopterans in South 
Australia, with a description of a new species of Bracon 
(Hymenoptera: Braconidae) - - - - - - - - 131 

Bird, A. F. Studies on the soil-inhabiting tardigrade, Macrobiotus cf. 

pseudohufelandi, from South Australia - - - - - - 147 

Kolesik, P. Rhopalomyia goodeniae, a new species of Cecidomyiidae 

(Diptera) damaging Goodenia lunata (Goodeniaceae) in inland 
Australia ----- ------ 155 

Nicholas, W. L. Robustnema fosteri sp. nov., gen. nov. (Xyalidae, Monhysterida, 

Nematoda), a common nematode of mangrove mudflats in Australia 161 

Smales, L. R. A redescription of Aspersentis zanchlorkynchi (Johnston & Best, 

1937) comb. nov. (Heteracanthocephalidae: Acanthocephala) - - 167 

Brief Communications: 

Baker, G. H. Seasonal activity of the earthworm, Gemascolex lateralis 

(Megascolecidae), in a Eucalyptus woodland in South Australia - 173 
O'Callaghan, M. G. & Beveridge, I. Gastro-intestinal parasites of feral cats in the 

Northern Territory - - - - - - - - -175 

Terrace, T. E. & Baker, G. H. Predation of earthworms by the land planarian, 

Australoplana sanguinea (Moseley) var. alba (Dendy) sensu Jones, 

1981 (Tricladida: Geoplanidae) - ----- - 177 

lyier, M. J. & Williams, C. R. Mass frog mortality at two localities in South Australia 179 



PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS 

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 






TRANSACTIONS OF THE 



ROYAL SOCIETY 
OF SOUTH AUSTRALIA 



INCORPORATED 



VOL. 120, PART 3 



THE BREEDING BIOLOGY OF LITORIA SUBGLANDULOSA AND 

L. CITROPA (ANURA: HYLIDAE), AND A RE-EVALUATION OF 

THEIR GEOGRAPHIC DISTRIBUTION 



By Marion Anstis* & Murray J. LiTTLEJOHNf 



Summary 

Anstis, M. & Littlejohn, M. J. (1996) The breeding biology of Litoria subglandulosa 

and L. citropa (Anura: Hylidae), and a re-evaluation of their geographic distribution. 

Trans. R. Soc. S. Aust. 120(3), 83-99, 29 November, 1996. 

The known range of Litoria subglandulosa is extended and that of L. citropa is 

revised. Population trends observed at the type locality during the 1960s-70s and 

1990s are compared. 

The advertisement call, adult colouration in life, behaviour and embryological 

development of L. subglandulosa are described and compared with those of L. 

citropa. The single egg mass of L. subglandulosa shows adaptation to the lotic 

environment, being compact and strongly adherent. The embryos and larval stages of 

the two species are very similar in shape and colour in life from stage 17 onwards, but 

are readily distinguishable by mouthparts. Comparative notes on larval behaviour are 

given. 

Key Words: Litoria subglandulosa, Litoria citropa, distribution, population trends, 

advertisement calls, oviposition, embryology, larval behaviour. 



th<n\n<tnm\ of tin- Hajwi Susieh "{ 5 rtWJ I WUbbh 120( T. K3-w 

THE BREEDING BIOLOGY OF UTORIA SVBGLAISDVLOSA AND L CITROl'A 
(ANURA: HYLJDAE), AND A RE-EVALUATION OF THEIR GEOGRAPHIC 

DISTRIBUTION 

by Makion Anstis* & Murray .1. LrrrudoiiN' 



Summary 

Awns, M $ | iTTUyOH!^ M .1, | |o%) The breeding bwtogj v\' lJt»riu .stth,t>lwuli<h>.\u and /, 
liyltdac). and u re-BYMluitfinn ol [heir geographic distribution. 7Whm. A' Sin. S 
November, 14%. 



itroffi) t Anur;i; 



nil 



'Mil- known mnge "| UWhti whxluitduiost; is cvlcnJu'J and lluu 111 /.. rmvyvj is ivvised. Population u 
observed at the type localiiy during ihc 1960s 70s and iy$ta flit! compared. 

;dl. adult colourafii'ii in lilR. behaviour and enihryological development ol / 
ibed and compared with ihose ol L. iitm/tu. The single e^'g mass ol /. \ithuhnulitii>s< 



I he advertisement call 
SUbxfanMo.Ht are devcriL. 

shows adaptation hi die loiic emironment. btfiruj compact and strongly adherent Flic embryos .ind larv.d slaves 
o! the two species are very similar in shape and colour in life from KtUge 17 onwards, hW $t£ rradilv 
distinguishable- by moulhparts. CompataMvc lotcs do lOj'Vat hctwvuuir are given. 



Kiv Wwkus: I if/nut stihtihaulttfosa, Utwitt cfflvpjtl, disiribution, population ireiidv. advcniscnieiii calls, 
oviposiiion. entbryohH/y, larval behaviour. 



Introduction 

Lttorio \Nhi>lit'itlulo\u was described as Lltv)'hl 
xlun(iut<Ksa Tyler & Anslis. 1*9?? but renamed 
because of primary homnnyrny (Tyler & Ansiis. 
1983% A member of cite L it'tropn species group 
(Tyler & Duvics [078), L suhi>latuitihi\d was 
previously Known only loan the Quecnsraiiti/NSVV 
border south 10 the New raigland ranges ol" northern 
NSW (Tyler & Ansiis 1975) The type description 
included a description ol the larvae, but no data were 
available on ovipositiomcrnbryoloijicnl development, 
larval hehavioni or the* advertisement cull. 

The species was found 160 kin soulh ol its 
previous known distribution in die mid-norih coastal 
ijm^cs and Burriivjton Tops region by one of US 
iM.A.i. in 1977 lis presence there din\ ihe absence 
ot / . tny>i>u, prompted a re-examination o\ the 
distnbubon of both species. In addition, observations 
on ovipositioiu the morphology <T embryos. )ar\ae 
and adults and a comparison of the advertisemcni 
calls i.i f L. ritn'pa and /.. sidiyjttndtdasn were ma<le 
and are repotted here. 

Materials and Method* 

Liu ft fa \ifh's>i(tti(lnl('\<t 

\didt specimens examined: Australian Museum 
(AM) RI7577. 35525, 4293i-33. 50I6.V 510&&-7, 
51104, 517354V. Point Lookout; R3445N - 14km 
f-.asi ol [dim: K36724 - Oafesv Creek near Poor, 



KWhtevrm Ril BetcrWRi Hv.yhls NSW 2oH2. 

l)r|iiiMiiuiii ul /...iL^y. t 'hH' i-,Hv ,.t WUiMi.iH. 
P.HkvillrVK- KJSJ 



R^6 l )75 - (my f-awkes Rivei, Cbor: R7I 109 -71 I 14 
Back Creek (Rarwick Riven ncai Point Look<nu 
R370I7 - 5kni S ol Walchu : R3O05C) 50km F of 
Cdeii Innes (Gibraltar Ranue); R5293I Samlw 
Cieek, Doiriijo, R5II7X-XO Sts \ River. Point 
Lookout; R76S|g - GlouccsJer Tops: RM6K3 
L Ipper Allyn River. Fiarnivjion lops; fcUMSOS 
Hllenborou^li River. Bul^a State Foresi, NSW. 

Lttnria enount 

Adult v/vr/wevM csaminctl: Atislralian Museum 
R75bO_ Orbi3st:75^ AbertekkVic: 19237. 18234. 
Uttfe. IK23X Slanwcll Tops: 794^0 Sianwell Park: 
245t»0-24505 T 2759t», l-mlc.mhridpe: 45X5X. 
rhillmere Lakes; 3I6S5. 71 12. 7S^>27 1 leiensburch; 
45424. Tianjara l-'alls; 5 I NS. Megaloue Valley: 7 1 10, 
lla/elbrook: SHHL Blaekheallv. 69034. Bell. 
Kurfifiuns Rd.: 76625. IS km N iit l.ilhgow; S459, 
Pennant Hilts; 14495, Colo Vale: 79100. 76623, 
Clljoul Range N of Colo flt>.: 4261. Bundainjon; 
7IK9S. 24 km N of Moss Vale: 15462. Goslnrd; 
/S264-26. 7S698. Kurmgai^Chase: 6<»425. Nad^ree 
Reserve; 794*9, (Jalslon doi-ge; 7563, Manly, NSW. 
Three adults iitcd as /. vitrofiu b> lyler & Ansiis 
(1975) irom Barri nylon Tops localities- l)epl/ool_, 
UtliV. Melbourne (MC/I)| 1792/64 - t'].»pei Allyu 
and MV/O 1690^1/63 - Wombat Creek, were re- 
examined because of appareni overlap in ranjjc with 
Ihc Barrirtrton Tops localities lor A_ vuhithmdidtna. 
These specimens have since been registered by the 
National Museum ttf Victoria (NMVt as D32666 
(tipper Allyn Riyei) and 03260T65 (Wombat 
Creek I- Similarly, two specimens <NMV D670'J- 10). 
I'iied by Copland (1957) i\$ I. tftfQpn from nr.n 
( irafton. iMMtheaslern NSW. wore examined, 



84 



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ft 



Q\ i/>o\itfon and rmhty<n 

Observations on three captive breeding pahs ol 
i-;ii li species, collected by MA-, art* summarised in 
lahlcs 1 and ?, In each case-, a caUin*i male was first 
collected al night, then a gravid female was found 
durin;: d oylight the next day. in (lie same vicinity aa 
the male The pans were each placed i^ a large 
ipfl&icd ptolU b&g ejOnftU&iflg siieain water, fi flat 
ruck and aquatic vegetation. The bag was covered 
Willi opaque material lor the duration of amplcxus. 

(Jtttriti \iih\;ktntlitlo\u 

SlftgCS |*23 (Gosncr, (9601 were studied ffOW 
three separate £gg masses one from (lie type locality 
(locality 10. Table I i and die others from \nc new 
localities 3 dlld 4b Hereafter, numhered localities 
will refer lo "fable I (unless otherwise stated*. 
plirthci '-.aiupte 9 froill eeg masses found ill (lie stream 
al locality # were maintained until stage 25 to 
confirm idcniitv. b\ I> A. White ot the National 
Patks & Wildlife Service. NSW (NP.VWS). I'mbryos 
and larvae Were held in dishes (40 cm dram.) 
i 'oni.iiMimj Mui.iiM Wiik-r. toekv sediment^ and 
aquatic vegetation, and maintained al 14 21 C 
(locality 9*. and 15 -24 C (localities 3 and 4b). 

Phc egg nwws TOW locality 4b laid on 7 ai i. 1 '/Htl 
< lahle J i. was submerged wilhm a metal lea strainer 
in the tool, flowing w.iter*.)' the stream I'm the initial 
two days ol development, but bolh Ihe egg masses 
from localities 3 and 4b were maintained at higher 
temperatures of up to 24"C away Irom the stream 
from the third day alter deposition. hmhryoiOL 
development was observed under a Wild \!J 
stereoscopic microscope, 

Liioriit ettnuiu 

Stages I 25 were studied from (wo egg masses 
Iron i Darkes forest and one from Ourimbah 
(localities I? & 16, Table 3). Samples of eggs found 
fullered over the substrate in Ihe stream were raised 
to stage 25 io confirm identity. Adults in breeding 
condiliun were placed in an inflated plastic hag 
covered with opaque material during amplcxus and 
lUcic^uUinc embryos maintained al lb -23 C\ 

Liirvar 

tadpoles were measured (lo 0.1 mm) with vernier 
callipers and an ocular micrometer attached lo the 
microscope They were anaesthetised in Chlorbutol 
solution before preservation in 37/ lotntuhn. The 
slaving system is that of Gosncr (IVoOl. 
Abbreviations for larval measurements shown in 
Table 0. follow Anslis (1976): I'L - total length, BL 
= body length. BD - maximum body depth, TD = 
maximum tail depth. TM = tad musculature depth 
(measured in line with TD). IO = inierorbital span. 
IN = intcrnarial span. LN - the distance between eye 



and naris and MW = maximum mouth width 
Illustrations were made using a drawing tube 
attached US the microscope. Preserved and living 
larvae ol /„ \\th^landuh>su fmtn sites I u were 
examined for comparison with those Irom Ihe type 
b h mIh\ and measurement* are given m Table 6. 
("ceding and swimming behaviour of several larvae 
of boih speuesw.r-obsefved m captivity uud in Mien 
natural Mfc environmenl. 

Advents nwtu nd(\ 

'I lie calls off. \tih^luruhiit>Mi were recorded at a 
lapc speed of 4 7fruiiMf.c "'. usiu^ a Sons Tt-ttSfRO 
portable cassette rccordei with a L her M5I6 
microphone and a Grampian paia6olu lerlivtoi. 
Calls of /.. L'hr&p& were recorded with a Nagra 4 2 
open reel lape recorder al <t tape speed ot |4 cm m\ 
and a Beyer M-XN eardioid dynamic microphone. 

for L, suh^lunduh'SOy the tape cassette was 
replayed on a Nuknnuchi Dragon kipe deck, and lot 
/., i ///r//w. the open-reel tape was replayed on either 

attevOVB 77 or uSo.n ■'!(. S I0O l;ipe nxoidei. the 

calls ^rffi analysed Oil a Kay r:lcmelnes Digital 
Soiiii -Graph. Model DSP-5^00, Addil tonal analyses 
ol waveforms were made by way ol a Sound- Blaster 
16 caid (Creative Technology! installed in an IRM 
K -compatible desktop computer, and using the 
Wiivv Studio (Creative Technology > and Spectra Plus 
Professional. Release 3.0 (Pioneer Hill) software. 
Both systems yielded consistent results tor analysis 
of the same signals. 

Ihe dominant (= peak) frei|uencu -\. wetv 
calculated as those ol greatest amplitude in a power 
spectrum or an averaged spectral display. Numbers 
of pulses were determined by inspection ol 
waveforms. Pulse mlcs were calculated from the 
interval between Ihe peak of (he first pulse and the 
peak ol the last pulse in a pulse tram and the number 
of pulses reduced by ona tie, n-1 pulses) Because 
of the difficulty in determining ihe beginnings and 
ends (i.e.. zero amplitudes! of pulses and pulse 
trains, the peak - pc;tk interval was taken as the 
duration. Where appropriate, pulse trains arc termed 
'notes'. II two distinctly different types of temporal 
unit are present in a call, then Ihe signal is described 
a* diphasic isensit Uttlejohn Sf. Harrison 19851, 

Results 

Di>lrdvitton und hahitui 
LUoriu snlygjitndnlosn 

The new loeahues { I -9) recorded in Table I extend 
the known southern range of this species about I.H0 
km All localities are permanent streams/rivers oi 
basalt or metamorphic rock country associated with 
rainforest, montane or wei selerophyll forest (except 
for Ma iSc Mb) anil are al S10 m or hicher. The 



go 



M AKSTISA M I I IT! I PJOIIN 




O — L. citropa 

♦ *=ti MibgtaiKfulosa 



100 200 



I ir. I. A revision ol ' (he distribution <>) ' I ttorut < '7//tyW stud 
luenu ufbglamhiUw provided by Tyler & Artttrh 
i l<W5k incHnhny ;t number of ne% Inc;i1ilux lur/. fcufa*. 

iatiflufo.sa. 



southernmost locfiljl) .il Wbicll the $pCCh$& has been 
lound is locality c >. Kal Brook, Mounl Royal Slate 
Porvst. NSW. Hie National Parks & Wildlife North- 
east Forests Biodiversity Study ( 1 091 I'ftH) riscprds 
/.. siibgUineiuinsa at a number of sites between ihe 
Harrington Tops region and Ihe northernmost forests 

«»i nsvv, including DoyJcs Rjvftr state Forest, mi 

Boss Suite Poresl, Nowciidoe, Wcrnkinihc National 
Park Gibraltar Range National Park, Sty \ Stale 
Pnrcst Spirnho State Forest and Boonoo State 
Forest. Phis indicates the species has a fairly 
continuous distribution along die range country, from 
locality 9 in the south lo near Slanthorpe. jusl north 
.,( the Old/NSW border (151 40'30"h, 2S'40'20" 
SMl-ip. 1). 

On a daytime visit to localities 10 M-lTu I I on 
1 9 \h.I994. no tadpoles of Ibis or olhcr species wcic 
located. This was at a tune when numerous tadpoles 
of L. MfhghHNfftln.stt, L~ h{)f>moh>m;fnsis and 
MtMipliyc.s hitlhus would he expected !o be present 
(based on annual studies in (be 1960s and '70s). 



Observations by John de Bavay and Paul Webber 
confirm thai (here has been little evidence of this frog 
over recent years at the type locality, suggesting lhat 
the species may be undergoing a decline there. The 
National Parks and Wildlile Biodiversity Study has 
records ol live nudes of Ihis species calling at three 
sites on 2.i\.l995 in the Styx River Stale Poresl in 
(he region of the l\pc loealiiy: 

I j Holy Creek -"lat./long. 30 _W 39" K, 152 14' 
4 VS. (altitude 1 (Kid ml 

2> h'elv Creek - 31]' 35' 26" b" 152 13' IK" vS, 
(890 m) 

3) Watile I U Camping Area - 30" }5 ' IS" P, \?1 
l2'3X"S(X70m). 

Observations on 20,\ii.|994 at localities 12. 13b 
and 14 (all northern localities), indicated the 
presence ol L. \nb^iaiuin(nsa tadpoles 



Litoviti i iiVDpa 

Specimens NMV D33&66 fl .'ppor Allyn Riven and 
FH2664-65 (Wombat Cicek) weie examined ami. on 
die basis (4* the indisiincl tympanum, promineni 
supral) inpanic lold and head width, were lound tci be 
L Hiih\>}ut)<luit>Kd- NMV 1)6709- 10 cited by Copland 
1 1957) as ( . < ifrofHi from near Oration in the north- 
easl coasl of NSW. form (he basis of ihe slalemenl by 
Heaiwnle ti uL (1995) (hat L vflrdfHi "extends from 
norlhcasicrn New Souih Wales (o southeastern 
Victoria". Upon examination, these specimens were 
huuid l»' have (he body proportion Ctf / 
suhi>Uui(/ulifsa t but because both were collected m 
1X65 and in a po<»r state of preservation, it is diffieuh 
in come lo a dellniie conclusion as to their identity. 
The iwo species have not been found in syiupauy ai 
any site examined, and this fact, in coihhiiiation with 
the examination o\ museum material, indicates Mui 
the drainaec o( the Hunter River appears to be a 
natural geographic barriei separating (hem iFie.l).. 

Larvae were observed by MA. on Pi. 1976 and 
25. i. 1996 at Boardinghonsc Dam in the Watugan 
Slate Poresl, south ol the Hunter iivei NSW (33 00' 
01" H. 151 24'l5\S)and by R.Wells lurlhcruoiih 
in the Pokolbin Stale Forest, near Cessnock.. in 
January 1993. Ibis fc ihe noiihcrnmosi known 
locality lor this species. 

Lttorid stihgUimlulosa appears to replace /.. citro/ut 
in the Bamuglon lops region north of Newcastle 
(lig- 1)_ /.. tirrttfhi occupies a wider variety of 
huhitau than L, .sttbxhitulalosa, including permanent 
streams in basalt country associated with wet 
sclerophyll or rnontane foresl, to similar streams in 
sandstone country. Although found at an altitude of 
1066 m at Abertcldy, Vic and Rlnekhemfv NSW. /, 
iinttpd also bas been lound m lower coastal areas lo 
50 m (locality 16, Table 3>. 



BKI-LDINC UIOLOliV Ol 



t'lUUANW'iOW A I., t </hy>r\ 



'•I 



Atltili vchmr in life 
I itnriti sah^hintlulesa 

Specimens from northern localities werti 
predominantly grefiji. whcicus those from mid -multi 
i <>,isi;tl localities ( I 9) ranged I'rom uniform golden 
biown with sc altered darker im tilling o\c\' the 
dorsum, lit specimens vvilh some small areas ol 
given often alone- the canthus rosiralis or under the 
eye. Two males liom hiealiiy I each had ;i broad 
dorsal paich ol i-jven over ihe head or dorso lateral 
ledums Two specimens. AMR7ro M), fuan 
tiloiiecsler Tops NSW. and another observed b> II. 
limes (NKVAVSi al Fal Brook (locality S?K were 
umlotm bright gisctl, apart rfbm the ehurucU'ii-..u 
ijolden dor.so-laleral snipes. 

Some vuldemhrown specimens dcvclopc. large* 
hntihl green palehes over llie dorsum al nielii iS. 
Cow pt rs, coium.k The inner surfaces ol the hind 
limb and gTQiil area were uanslucenl yellow, as 
touud in adulls irom the type locality. 

l.iioriu titrofht ha.-v a uniform golden brown dorsal 
colouration Iwiib green along ihe canthus rosiralis 
and sides of the body i. similar to most specimens of 
/. sith^/atulnhtstt from localiliC-% 1-9. The principal 
difference belween the specie*, is the colour of the 
inrici surfaces ol The hind hmb and eroin, winch m /.. 
i iit.'fii is brick red. 

I ttftfHH >t<'tntt\ 
l.tlcnn mkglttitthifoiii 

Culling heems in spring and vvns observed on 
20a.I9 1 >4 at liualiiv X. when water icmperaiuies al 
IligllC were very low. e.:: . n C and the div bulb an 
temperature al locality 5b (1900 h) was 9,8 C i.A. 

While. S, Cow |vis_ LOinm.t, Olhei obsei VatioUs by 

M \. al ihe type locality during annual three-week 
periods (Dec/Jan. I9fi(i -74). and al all other 
localities listed in lable i. mdieale ihut calling 
persists ilirouidiout December/January in a variety of 
ur.iihcr conditions, with increased i.etiv ily during, or 
aflet. heht rum. livening dry bulb air temperatures 
taken during pcrn'ds ol spring/summer aclrvily al the 
localilies in Table I wete |V"C-I9.5~C (mean 
15 7 O At the lower temperatures (>V-I4"C). 
calling wa.s less intense and by aural comparison 
onlv, notes were al a slower repetition rate. 

S(M»iadic dunoal calling was common during the 
bteedme season bul males were most active at me, it. 
Diurnal calling look place from concealed positions 
such us under rocks or from within vegetation, either 
near the siieam. ot at time* up to abotil i'om meues 
away from the water. A single male or a small 
number oi individual-., called from as early as 0742 
h (e.g.. locality 7b). Nocturnal calling was initiated 
by one forjg, in-imal!*, followed by others m ;i 



distinctly polyphonic chorus The calls oi frogs at the 
souihcrn localilies could nt>( be diHercnliulcd from 
those of males ,u the lype localilv. 

Males observed calling al night were often petched 
on broad leaves of trees and shrubs approximately 
0, > 1 ,5 m above streams, on terns at the edge of the 
stream, or on vcgclulion lurlhcr from the waters 
edge. They were frequently found calling in small 
groups, iwo or moiv metres apart- On 22.\AW-\ at 
Inealiiy 5K. 40 males were catling at riigtil in groups 
•)f up to six alone; a 50 m >nvtcli of the stream iS. 
Ciow pers, eomm.». Al locali'y 7b on 7,\i.I^'M. lour 
males were ealline 25 m apart (K, Thumm |vr>. 
comm r 

An analysis ol the advertisement call is piostded 
below and comparison made with thai <d / fftm^i 
lw<> adiitliiuud call sequencfs. alliibutable 10 /.. 
stfh<;iiitutf>h>\ii y 3TC in the Btoaeouslic Library ol Ihe 
Depal'tutenl of Zoology VniNersKy of Melbourne, 
both recorded bv M. J. Liltlcjolui and bis associates. 
The first, from tiuy Kiwke> Creek hbor NSW i Mi 
24' 20" L. I.S2 20' 4h" Si \\<i\s a-emded on 
2JS.\, 1^64 al a wel bulb air temperature ot :<S C. and 
Ihe second, from Flat Rock Creek H km W t^'Poiiil 
Lookout NSW (close to ihe f&W silt), on I L\.1V63S 
ai a wet bulb ail lemperaiure of I VC They ate 
similar in all pertinent respect> to (he ljII ilcseriheo! 
here. 

Lt!<.-riii ci/ropa 

Males al Darkes horest (locality 15. /'able \\ were 
obsei^ed dunn» s[iriuy and summer catlm;j lo> t (j low 
branches neside Ihe stream, on rocks near the otj&e ol 
the ^.itei. iv on exposeil rock shelf in midstream 
clove to shallow, slowly flowing water \> \\n\\ I 
MibyJiittJiilt^tt nulcs called while two or more 
melres aiarr and activil\ mcicised on oVercasi 
evenings during or aflcr rain. Pry bulb air 
temperatures on several nighls when males were 
calling in Septembei LXxernbet. 1972- 1VS0. wete 
14 22 'C No diurnal calling vsa% obsetved. 

Aihcrfiu'iwtil *v(h 
l.ctena \nbitlutuhilosit 

Tlte .idvertisement call of /.. wfa$IiWfItfif}.st{ was 
recorded by .1. Courtney al Diehard Creek, Clen 
Innes (locality Hai. on 20.\»3. The dry-bulb au 
icmpcrature was I V C\ The lollowine, data were 
obtained from the fourth call ill Ihe sec)uence (I : \ii. 
^A). The call has a duration ol 9.375 s and consist,-, 
of 13 pairs (doublets! of pulse trains (notes), svilh 
each of iho.se in the first five pairs all beina of 
relatively low amplitude ibi", -mi. In the subsequent 
seven pans t)l noies. ihe second note is of much 
greaiet ampliiude than the firsi. Thus, all but one o\' 
(he firs! notes (which is of equal amplitude) are 
Softer ft 'I' 'he ani|4iliide of second notes bejf\£ 



l )2 



M ANSTIS & M. J. LITTLEJOHN 




Fig. 2. Live egg mass pf Litoria sub#lanchitt/.\u attached to a leaf from submerged overhanging foliage in Tuckers Creek, 
Burrington Tops ( locality X). Scale bar - 10mm. 



BREEDING BIOLOGY OF L, SUBGLANDULOSA & L. CITROPA 



93 



» hh |( 




8 



10 



seconds 




Fig. 3. Waveforms of advertisement calls of Lltoria sub- 
gkmdulosa and L. citropu. 

A. The complete advertisement call of Litoriu sub- 
glandulosa from which the values given in the text were 
derived. This call was recorded at Diehard Creek, Glen 
Innes, (locality 1 3a), at a dry-bulb air temperature of 
I3.0X. 

B. An expanded waveform of the eighth doublet in the 
call depicted in A. 

C. A waveform of the complete advertisement call of 
Litoria ciiropa from which the values given in the text 
were derived. This call was recorded at the Rocky Rivet- 
Road crossing on the Brodribb River, 17.5 km NNE of 
Orbost. Vic. at a wet-bulb air temperature of 17.5°C. 



100 



200 300 
milliseconds 



400 500 




1 2 

seconds 



*M 



M MXK'IIS Si M I I I I'll KIOH\ 



gieutei -by UpUi^dB in pair* 1 7. unci hy 12 ro 2*) tit*- 

in pairs 8-13. Durations of doublets ranee Irorn KM 
ta 372 nm (mean - 334); intervals between doublets 
liinv from K4 ro 154 nis (mean = UK). The 
ntpCti'Ottti rate t>f" the doublets is 1.35 s' The 
• i.-itHiuini oi peak bequencics are within Hie Mftgffri 
I fo(M480 H?i with means of 1405 flz lor tfe RrM 
notes and 145-1 H/ for (he second notes. Thcie ate 
10 15 pulses (mean =■ 14.1) in the fjrsi notes of each 
i»t' (lie first seven pairs, and 4-f> pulsed (mean - 4>:> 
in the remaindet. with 13-23 (mean = lf>.l) ill the 
second note of the lust seven pairs. and ''-1 1 uncm 
a 10.3) in iltc remainder. Ranges ol durations tit first 
rlilUb ii- J2-*)1 ins (mean = S5A>i lot ihe fust seVflfl 
pairs, 43-WJ ms (mean = 5*J.N) lor the others ;.uid I _'o 
I IV m I mean 137.61 mJ I U 15\> ms imeau H4.2> 
respectively loi the second notes. Pulse rates ot first 
nolo nmee bom 125 IA7 p s' (mean 152.5) in the 
tuM jyvcfl pairs, m 40-SS p s ' intern t\4 0> in the last 
m\ t-nr the second notes. |lta2 C&WgCi '"• fU)|sC fa 
lire l ' 1 -IX1 |? s ' Muean 145.01 for Ihe first ta en pairs 
and S6-75 p s ' (meiin b4A0 for the last sjv 

f.font, r/ffifpa 

the udverli%ciueiil call £rf (his species was 
uY.-anheil by l.uilcjohn t/ ^//. ( 1472) born the 
audiospectrographic and oscillographic analysis of 
two calls of one individual recorded (Nauru 1MB 
recorder, hkvlro- Voice r£V 644 microphone) M 
l.Hi.eh. C«cfc 9 km W uftann Rivet Vk (I I9' 1 OS 1 
I . ; . / 34S) on 24 v!'>o0. The male was calling on 
tin* bank al a wet bulb air temperature of 10.5 (\ 
Dwitfg lo background noise levels in the recording. 
only u tracing of" a waveform was provided 

This relatively long call (32 3.6 s) was described 
as *»l complex temporal structure l|j£n 4rougl> 
diphnsiek with a long introductory note lOM)- 1 ) 1 *) 
ms i erf. high and regular pulse tate t46 ps ), foil owed 
by a sequence ol inegukuly produced pulses id 
groups ot '5-7. The groups bave durations beLweeu K0 
and 120 ms and pulse rates ol' 34 57 p s ' \u_-a\- the 
stail. and are lottj#J (2^5-500 W$) and ot hn\«t pulse 
rale ( 10-21 s ') near the end. The dominant 
fivqucntics winge horn 1350 to IK0O 11/ within a 
broad band ol frequencies between 1250 to 3600 H/. 

lo confirm this desetiplion. and to provide an 
■ itdk'alion t>f 1'nissible effects of (cmpr-oHnrr, the laa 
cleat call in lire recorded sequence ol anothei 
individual ot I,, i ///?'/></ was analysed. The recording 
was made at the Rocky Rivet Road crossing on lite 
Bmdiihh River 1 7.5 kin NNL o\ Orhost Vic ( 14* 
JJ'E, 37 30* Si fay M. .1 Litllejohn on 2N.xi.l l )KI. 
This, frog was calling from vegetation ill a height of 
about 50 em, adjacent to the river, at ft wei bulb air 
temperature of 17 v C, A wave loim of this call is 
prcscnled in lag. 3C. 

Tlie call, which has an overall duration o( \Xbh \. 



ctHisists t)l a distinct lirsl note which is u iv^tilai 
pulse train with a duration of X14 ms. a pulse rale ol 
187 p s' and a dominant freqttencv of Ib40 H/ \ 
single pulse (duration appn>\inialely 7.0 nisi wjlli a 
dominant frequent. \ oi IMIO Hy follows The 
oiii.undei ot' (fie Call consists Ol eleven ^ioUj^ ■-! I 
5 pulses but four pulses and one pan ol pulses eante »' 
rcalisticallv be grouped to alknv calculation of a 
pulse rate: otherwise pulse rales ran^e from 31 to fU 
p •-, '.The di.niiiumt fret|Ui:itcii:--. ot 'these pulse, OMtp^ 
from 1200 lo I3'M) 11/ and ibe durations ot the pulses 
ranee from S.6 lo 12 9 ms, The variable pulsatile 
second p;u( ol (bis call has a maximum amplitude 
about ^ i||"j higher than dial of the iiiltodueiory note 

Liietia whglattduhtsu 

The adverttsemetil call of the male was heatd ill the 
hffg before an. plexus occurred. Details ol egg mas -c. 
laid aie pieseuted m laMe 2. O\qiosi(ion was not 
observed, but lot each ol the three captive pairs 
sludied, a single e^g mass was louud adherine, to the 
side o! die bay..|Usl below walet level. I he UggS were 
laid in a small, cianpaet clump ol 1 iwo lo three layers 
id extremely sticky, coherme capsules. If. 1 
Loniplements Tor two Ivmitles were 2V2 arul 4_\ 

Another eight egy masses o\ [ftis species were 
louud at locality S on 4 xi. I W4 by A. While and 
S, Cow. Hach mass wa.-- auached t« an oveOtaugiiig 
leal a tvug oi a ovk [tyM U) vertical 0! neat \co>- -..l 
orieulalion. just below the \\u\l'\ surface in a slowl) 
llowing section ol j pool (some in mid-stream), The 
pool was heavdy shaded by an almosi compiri: 
l aio[*y I'over. Steadv lam had fallen Ihree ilaj 
eaihcr L .od the stirijice water temperature a# 
1400 h was 'J 4 i \ One o| ihese masses, removed 
Irom Ihe stream on a leaf and photographed, >v 
shown in fig 2 

Lih>iiti ittn/fitJ 

On 25-vi-l u 7.V oviposdion occurred alter (he m. ! 
t i\\i\ teiitalc had been collected at Ouriuibah ("reck 
NSW (h^calily 16. Table U ;il 2200 h. The Imps 
were placed in a plasiic bag. I he male soon began to 
call and the pair was m amplexus ihree hours abet 
capture. At 015b h on 2o.\i.l < >73. the initial two 
sequences ot'o\ iposiiiou activlls ocetined; a( 0157 h, 
a further lour oviposit ion sequences followed. Willi 
only about three seconds between each. Oviposidou 
was complete by Ol5V> h 

In a typical sequence the female dorsiflexcd her 
body with otilstretched hind limbs and produced a 
hatch of eggs. The male fendistd them while 
clipping his feel in a tanning motion around the tggs 
The leiualc then seaUcred (be eggs with three sudden 
k ictsine movements of her hind limits, The egg.s sank 
and spread in a single layer across ihc bottom. She 



BREEDING BIOI OC.V Of I, SUBULANWl OS\ ft /- C77W/M 



U5 



T'AHI.1 4, < 'itmptirixfln of r emhr\t>\ ^/'Litoria suhglamliilnsa flnrf I Unria eit opa 
Developmental stages are those of Gosner ( I960) 



Sliitii 



17 

21-22 
21-25 



Sam pi 


e 


Mean emhivo 


Mean capsule Description 








diameter/length 


diameter (mm) /- Mtbghimiufwto 


/ , ( tlli'f'tl 


h.Mih^lnntL I 


..iittvpti 




( mm t 









D / 


Mf/U,'/ 


tW. /.. Litrt'fto 


L subgtand, /.. tfmopd 




•J 


» 


1.59 


I..73 


3.5? 6JTJ Animal pole: black/ 

vi'Mi'ttit />"/<': dark grey 


daik brown/ 
creamy while 


g 


1? 


2.17 


j,!2 


Body: dark grey, 
yolk sac light grey 

//ti«/ (lalciul view): 
acutely angled 
tyffr ve.Mi/c; small, 
distinct bulge 


dark brown/ 
ereamy while 
acutely angled 

larger, 
indistinct bulge 


1 





653 


d.76 


Gfte anterior 3-4 branches 
posterior 4-5 brunches 


1 -2 branches 
2-4 hraiuhes 
(shurter) 


1 


8 


Z.39 


u.05 


Uttered lines: pigmented 
Mtxtfh-parys; no looth town or 
keralimst-d |a\v sliealh 

H(jW'liiri& U'mt't 0-10 day* 


non-pigmenled 
luuih rows, 
keralinised 
jaw shealh 
4-0 davs 



ihcn swam 10 another &kc and the process was 
repealed. During ihe Final sequence, the emale 
remained in the dorsillexcd position about three 
seconds Longer, bin produced no eggs. I he male ihen 
released the Female at the poinl when she began 
kicking her hind limbs. 

The egg complement was 055. F.mbryos hatchet! in 
tour - five days at waler lemperalures o\' 17' 23 C 
t\vw other egg masses laid m captivity contained N00 
eggs and 928 eggs and look Tour - six days to 
complete lunching at In -21 C (see Tables 3 $ 4). 

In llie field, ihe eggs were found scattered 0VW the 
substrate HI shallow pools or slowly flowing sections 
6f the stream, which is similar lo the mode pi' 
deposition of eggs observed ill captive pairs. 

t \t<i i 5. ninwiisions nj prewired emhryos of Litoria sub 
glandulosa 

imeaji in mm. range in parenthesis, stage 
Gosner, l%0) 



S la i:e 


Sample 
4 


l\mbryo diam. 
1.59 


Capsule diam. 


i 


3.37 






( 1.30 1.04) 


l3 12-1.5! i 


7-X 


5 


I.sfi 


3 50 






(J .56 1.30) 


1 ; IM.85) 


16 


s 


2M5 


S.3? 






12.03 2.03) 


1 1 2S 3.53i 


17 


9 


217 
i2.M-2.Ui 




20 


fi 


5 SO 
1 5 49-0.23) 




M 22 


\ 


io* 

(6.4N-0.O4) 




23 


1 


0.80, 7.3-4 




24 


1 


7 V) 





kmbnontt (Icvclttpmcnt 
Liior'nt sttl>#hiiulul<tsa 

Mortality rates ol embryos maintained in captivity 
were high. Tilt survival rate taller removal from Ihe 
stream), was greatest amongst embryos in the lop 
layer of each mass. Those below this layer mostly 
ceased developing beyond aboul stages K-12. 
Kmbryos from the egg mass held at locality 10 m 
water temperatures oi I4 ff -21°G survived the 
longest; hatching occurred from days S - 10 and only 
17 reached stages 20 - 25. The mass from locality 3 
did not develop beyond stage IS, Initially, Ihe 
embryos from locality 4b continued lo develop 
during the two days of immersion in Ihe siream 
before higher lemperalures away Ironi the stream 
were experienced. Hatching occurred at stages 20 - 
21 from days 6-8. with only eight embryos sui 'vi\ ing. 
Embryos from the southern localities match the 
following description of those from Ihe tvpe locality 

10. 

Imibryos laid early on I. i. 1974 (locality 10) were 
at stage 2 when a sample was preserved at 0945 h. 
The animal pole is black and the vegetal pole dark 
grey. There are two layers ol 'jelly surrounding the 
perivilelline membrane. Measurements of embryos 
are given in Table 5. Ihe embryos were at stages 7-X 
after K h. and X-9. after 12 h. Six embryos at Mages 
7-5}: measured after preservation, have a smaller 
mean diameter than Ihe same embryos measured live 
(1.7 mm live, 1.5 mm preserved; capsule dunnctci 
3.0 mm live, 3.3 mm preserved). After 23 h. embryos 
were at stages 10-11, and after 38 h. at stages 12 H 

Stage 1 7 w as reached after 02 h. A specimen drawn 
at Stage 17 (Fig. 4Al. is described:- prominent optic 
vesicle, pronephric swelling, slight anal bulge, large 



% 



M ANST1S& M, I- LITTKKJOtlN 













B 








' i ■ i . fnfifjrViiS tif t ihnhi \uh^lamlnh>\n ;m*t /- i >tn>pa 
\ I- \ithv!<in,Jiihi\,} rv'inmoil Irotii its capsule al Maiv 17 
li / i Hi'tpfi ivriM'VcO rprtm its capsule, al Ma^c 17. 
t / -■iihvjuthiulo.ui jutl hutched, at approximately *tyg£ 21 
I) / Ultiipa fUtt lunched, at appiuMiiukU Muyc 21. 
S. :,!<• bllt * I mm Ma'.v. hit I'rvm I • ■ n: a I I '■ 



UKLliDINCi UIOUXiY <)1 / St'HdiAMHtDSAX- I I V/A'M/M 



W1 



gill-plaic swelling, with beginnings ol muscular 
ridges along dorsal surface just below neural tuhe. I' 
shaped adhesive organ, slight stoniodueal groove 
beginning io form. Head iruneaie. acutely angled in 
lateral view. Tad bud short, rounded, with strong 
depression on each side below neural lube. Voik sac 
grey, rusl of body very dark grey Aitei some years in 
preservation, hody appears dark nod yolk siu hghtet 
brown. 

limbiyos examined at 71 h were in stages 17-1 S:- 
growing tail bud pointing acutely to the left side of 
I he bod_v within rum jelly capsule- two visceral 
arches forming, narial pits beginning to develop 

Alter 95 h. siaee IK-- optic vesicle more dellned 
witli groove forming between this and gill plate; 
neural tube, dorsal muscular ridges, narial pits and 
divided adhesive organs all more developed. 

Aftci 13! h Mages 19-20: small external gills, gill 
emulation not apparent; head small, more rounded 
over cranial region, adhesive organs dimimslung; 
optic vesicle depressed slightly in centre: live 
cmbiyos dark grey dorsally, lighter grey over yolk 
sac, moving actively within eapsiile. 

Hutching began eight days after ovipoMhoti- all 
surviv i ng cmbryoN had hutched after ten days. 
I mbryos hatching liisi mi day S wcje at stage 21 (in 
relation io nplif development, but io gill 
cireulalion):- Optic vesicles indistinct, yolk ?HC 
large deepened slomodaeal pit with adhesive organs 
close loi'ether al aiiterioi end, a divided ridge 02 
posterior end; gills, developing notfeeablv UnoIX 
advanced on sinistral side: vcnl tube not well 
diflerenlialed: tail I ins dusky grey, shghdv arched 
dorsally; body dalk gtey l>imvn m pre-ervati\c. head 
rceion dichlls darker. 

Stage 2 I was reaehed on day HI in relation io gill 
(leMiopuient and lack ol tail lln circulanon only. 
(big. 4C)r- iwo pairs ol well developed functional 
external gills, composing 2-1 branches on anterior 
pan. 15 on posterior pair; adhesive organs small 
translucent; opli»_ vesicle undefined. Tins translucent. 
deepening further, circulation not apparent; iaii 
musculature poorly developed, 

hive final hnlchlirigs at 19(H) h on day 10 were at 
siaee 22 in relation io tail circulation, hut other 
development was associated with stages 20-21:- 
corncu still not irauspareni, prominent bin only 
partially pigmented optic vesicle, lad fins deepening; 
gills al maximum development, fully functional, 
longer in some specimens than others: adhesive- 
organs merging to form small ridge*, mouth 
triangular, line ol pigment from lip of snout through 
each initial pil to eye. 

Stage 2-1- cornea transparent, eyes well developed, 
heauly pigmeuled; anterior hall ol body becoming 
transparent around nares; gills diminishing, operculum 
developing. 



Slage 24.- venl lube more discernible, oral disc 
developing, with small triangular funnel above large 
oval depression lo become lower labium. 

By day I 3. most remaining embryos were at early 
sta^e 25:- golden iridophores si/atteied in >pois ovci 
dorsum. eyes black wilh scattered golden indophores. 
patches of melanin over dorsal surface of tail 
riiuseuktiuie; lad fms. hody wall mostly clear, wilh 
some dusky pigment present Iniernarial region 
inuiceahly delineated with pigment, lateral line 
organs becoming visible. 

By day 17, the development of the mouth was 
almost complete wilh the exception of the line black 
filameuts, winch were either not yet present, or only 
shon unpigniented mots. Dorsal surface lurthci 
pigmented wilh more golden iridophores over ateas 
pigmented with melanin, including iris; tail 
musculature pigmented dorsally, in well-spaced 
broad hands: flecks of pigment found over bus in 
older iarvae. as >el noi obvious; ventral (surface cleat. 
cxcvpl lor broad perimeter of iridnphoies. 

Lit* >t la lim.'pn 

[jnbryonic development was described by Kin 
& Anstis ( l^7Sv. A comparative summary of embryos 
el /.. tthf'pa and /.. siih'<l(iiulnh>.\o during stages 2. 
17, 21 and 25 is given in Tahlc 4. Figures 4R, 
D show singes 17 ;uid 21. In general. L fiiprtptt is 
larger than /. vittykaidlilttiW ihmuehou! embryonic 
development, with adhesive oignns rt|tiic pioimnem 
and eiils smaller and less numerous at stacc 21, A( 
stage 25 and beyond, ihe lateral line organs remain 
iinpigmemed :ind mouthpaUs possess tooih rows and 
a keraiinised javv -heath tfig 4. Tyler & Ansiis 
1975). Otherwise, the iwo species hove distinctly 
truncate, angular heads in stage 17 and similar 
hotly/tail shape throuehoui embryonic and larval 
development. 

Ldn'it! htltintont 

Tadpoles of this species- observed at all localities 
in Table I. were mostly foiiiKl on the subsiruic in 
shallow, stovvly-tlowiug seenons of the stream nu 
sand, amongst tocks or leaf liner. They weic 
fret|ueulty found at die sides of the stream, 
.swimming fast to deeper mid-sireum or amonesl 
rocks it disturbed, They were well camouflaged 
whilst on sand or grazing amongst rocks and 
appeared lo feed on flocculem silt and algae. 
Tadpoles defaceaied nipidly after capture and ihe 
abdominal region, while similar in width io ihe 
branchial region fov slightly less) m liu- specimens 
in Ihe siream, was commonly iiatiovver in preserved 
specimens. 

Tadpoles observed adhering to die subsume 
rapidly palled the body forward a distance ot 1-3 mm 



98 



M. ANST1S & M. J. LITTLEJOHN 



by the use of the oral disc alone, in a rasping action. 
This process was repeated continually, resulting in a 
distinctive form of locomotion during feeding, which 
has not been described in other Australian suctorial 
species. Particles of a fine silt suspension were found 
amongst the dense, incurved papillae, buccal cavity 
and gut of recently-captured specimens. 

The fine black filaments of the mouth were broken 
or missing in some specimens, or each was present 
only as a shorter white filament or core, without the 
black outer surface (or pigmentation I. 

Litoria citropa 

The tadpoles were found in small rock pools 
(either associated with the main stream or segregated 
when river levels were lower), and in larger pools or 
slowly flowing sections of the stream. They were 
also found on the substrate, but unlike L. 
subglandidosa. were not observed moving forward 
by the use of the mouth alone; the tail and body were 
also involved. They appeared to feed on flocculent 
silt and most individuals examined live in the 
streams, had well-filled intestines (the abdominal 
region being as wide as, or wider than the branchial 
region). When disturbed they took cover under rocks 
or leaf litter. They were well camouflaged on the 
sandy floor and the dorsal colour varied from light to 
darker golden brown, depending on the colour of the 
substrate and light intensity. 

Discussion 

Population trends 

Comparative field observations of the 1960s-70s 
and 1990s showed a marked decline in the 
population status of /,. subghmdulosa at the type 
locality, indicating a need for comprehensive studies 
on population trends oi' this species across its entire 
distribution. 

Advertisement calls 

The calls of Litoria subglandulosa and /,. citropa 
differ markedly in structure (Fig. 3A, C) and cannot 
be of any assistance in the confirmation i4 
relationships based on other criteria. As noted by 
Watson et ai. (1991), the audiospectrograms of the 
advertisement calls oi' L. citropa and L. spenceri are 
of similar diphasic structure; they differ, however, in 
that the following notes in the call of L. spenceri are 
more regularly pulsed and of higher pulse rate. 

Oviposition and embryos 

From observations of oviposition sites of Litoria 
verreauxii, L. dentata, />. phyllochroa, L, caerulca, L. 
chlon's, L. freycineti, Limnodynastes peronii. Lint, 
tasmaniensis. Lint, ornatus and other species of 
Australian frogs, it has been noted that each deposits 



Tari.Ii 6. Comparison of body proportions of larvae of 
Litoria subglandulosa. 

Type Locality 10 compared with new localities 
2, 6a & 7a (Table I ). 

(Measurements in mm; mean with range in brackets). 
Stages 35 & 36 (Gosner I960). 



Morphnmetric 


Tvpe Locality 10 


Localities 2. 6a, 7a 


Character 


n = 8 


n = 8 


TL 


29.84 


31.50 




(26.40-35.00) 


(28.50-33.75) 


BL 


12.19 


11.88 




(11.64-12.63) 


(10.82-13.13) 


BW 


7.42 


7.64 




(6.15-8.04) 


(7.05-8.45) 


BD 


6.17 


6.10 




(5.74-6.64) 


(5.58-6.72) 


Tl) 


5.86 


5.87 




(5.17-6.48) 


(5.42-6.40) 


TM 


2.01 


2.35 




(1.64-2.29) 


(1.89-2.71) 


IO 


2.49 


2.75 




(2.13-2.87) 


(2.46-3.29) 


IN 


1.88 


1.94 




(1.80-1.97) 


(1.80-2.05) 


EN 


1.46 


1.37 




(1.15-1.64) 


(1.15-1.64) 


MW 


4.55 


4.48 




(3.77-5.25) 


(4.10-5,00) 



eggs in a similar manner whether in the field or in 
captivity (Anslis 1976, Anstis, unpub.). Similarly. L. 
citropa scatter eggs over the substrate in both captive 
and field situations, and L. subglandulosa attach the 
entire egg mass to a surface just below water level. 
The egg mass of L. subglandulosa is adapted to the 
lotic environment, being compact in form and highly 
adherent. 

Embryos of L subghmdulosa that survived beyond 
stages 8-12 were mainly from the outside layer of 
capsules. Mortality may be attributed to reduced 
oxygen levels associated with higher still - water 
temperatures in the laboratory of up lo 24'C. 
compared with 9.4°-15 Q C in flowing streams. The 
embryos from the egg mass at locality 4b continued 
development during the initial two days of 
immersion in the stream but, alter removal and 
placement in the laboratory, development gradually 
ceased over the next four days in the majority oi' 
cases. 

The periods of 6-8 and 8-10 days taken by two egg 
masses to hatch (while maintained in containers) are 
slower than those of other known stream-dwelling 
hylids of lower altitudes, including L. citropa (Tyler 
& Anstis 1975; Anstis unpub.). Further comparisons 
can be made when data are available for 
developmental rates of egg masses within the stream. 

The egg capsules of/., citropa. are not as adherent 
as those of L subghmdulosa. As they are scattered 
over the bottom of still pools or very slowly flowing 
sections of the stream, stronger adhesive properties 



HRLUDlNCi uioi.oc.v Ol L WUCilANfrULOSA & /- CITHOf'A 



vm 



would not be advantageous. The embryos developed 
faster and had a much lower rale of mortality lh;m 
thpSC Of L yuhxlwi(ltthi\<t. possihlv atiribulahlc tO 
Ihc individual capsules being scattered over u broad 
area, facilitating ovytienaliou. 

ItirvtW 
Whilst alight dilieivnees m body pi'OpOtUona WCIC 

noted bclwccn some of Ihe norihern and southern 
tadpoles of L wbglatulnfosa (Table f»). only y small 
sample from each area was examined. 

A sample of £. \nhi*lan(Info\a ladpoles wax also 
very difficult to maintain in captivity at higher 
Icmpcralures and a second sample maintained in 
ueiatcd water with liltraliou tared no belter. Lucking 
kcraiiuised jaw sheaths, they could not eat foods 
such as boiled leiiuce and commercial fish food, 
Introduction ol silt and deltital sediments taken trout 
Ihe streams in their natural environment resulted in 
some feeding, although llie tadpoles did not eiow us 
well us those in the streams. 

ThC distinctive locomotive behaviour of the 
tadpoles imohme forward propulsion wilti Ihe use 
of the oral disc alone, distinguishes (herd from the 
sunilat sympatic species L, fthxtttnhrtia and / 
/(■Mfcttii. bold of which employ some tail movement 
during locomotion associated with feeding, Ciradwell 
( I *>7> > stales thul the M3c muscle in /., 
Mth^ttiiuhtlosa tadpoles is inserted in both Ihc tipper 
and lower labia, resulting in both labia being "pulled 



caudad simultaneously", whereas "most other 
suctorial ladpoles move their upper and lower jaws 
towurd each Other during their scraping action". T his 
could explain the mechanism behind the distinctive, 
movement observed in live ladpoles in the stream. 
Gradwell also notes that this species has. lor its si/e. 
"Ihe longest and densest papillae of the buccal 
mucosa", and these "may aci as a sieve to rsclude 
suspended particles above u certain si/.e", 

Examination of gui contents and lurihcr 
obscivalions ol (ceding mechanisms arc required to 
determine the funcdonul morphology of the uni(|ue 
moulhparls of ihis species. 

Acknowledgments 

Wc are gniiefu! to the Australian Museum and the 
Museum of Victoria for access lo and loan ol 
specimen.^. The NSW National Parks & Wildlife 
Service is acknowledged lor permission to relet to 
records of Lihuio whfthttyiultfXtr. John Courtney 
provided the tape recording of /. \ubxhmfaln\n 
analysed. Arthur While provided figure 4, and with 
Karen Thumm. Jaequie Kecsei and Ihe late Shane 
GOW, added valuable observations on I 
MihyJiUiJulosn lo Ihis study. David. Dulcie and Ron 
Anslis, John de Bavay and Roy Scott assisted wiih 
field work. Stephen Richards, Michael Mahony. John 
de Bavay, Fred Parker and Michael Tyler 
eonstruclivelv reviewed the manuscript, 



References 



Am vi i . M I I07hi HivediUu hiul»nj> and larval 
Jrwlopiiu-nl i>l / iU'iui \n>r<!tni l An lira llvlitt.iel 
hwis. A' Soc. S /Wo/. 100. t93 2132, 

( 'oi'i \M.. S. I. (I $53 J Australian hei- bugs ul tin lthii-, 

Hviti Hiw. tiuw tow vs\i J&.U] 'J-itw. 
i i< i^ni-.k. K L. I |960l A simplified table lot sluying uimran 
emhrvus .nut Imvne wiih notes Oil I dull Til eation. 
fidprK'I.H'tn, |6. Ntt-IWF. 

( IK Oavi i ) , \_ t \ , PS) l^pctiiiK-nls on oial suction und ;:tll 
breathing in live species of Australian tadpole 
U\imra:llvltdnc and I .epiodnctvjiduet, 7, AW., htmliW 
177. XI */x 

Mi .m woi i . M . in ow.v.,.1.. Wi IUft.1t. I J & Wi:nn.(i ( |W5j 
I -iiitit.il Survev of New Knejand. tV. "t'he ft'qg*. flfan Old 
Mas Mii\h 224-344 

Ltiiia jomn. M. j. t v IUki'isos. pa. t 0>ks> The functional 
sivntlicance ol ilk* diphasic advertisement call ol 
(i<i<*>t»ui \iiu>>i<t>«> (Aiiui.i I i-ptodaelvlidaf). HfhtlW 
I n,l mulSntiuhinl, 16, M.V373. 



. I oi ns Hills. I, .! . Maioin. A A K VVaison. 

i_i. I_ I 107?) Amphibian fauna of Vielona Conhnnalion 
ol records of Uforiti <-lfyht) cifn>fni Cfseliudi) in 
Uippriitnd \t<t \,</. s*>. 51 54, 

I VI k. M. J. & Avsii.s. M. i l°75i Tuxoiiotns mid hiolo^s 
tlj hops ol rlw* Lih'ini i ffRtyW umiplex i Aiuirallylidaei. 
A'-,. V \,<st Wtty I7(5),.)|-M). 

A. tl u S.-'i krplaccinenl nunu lo. 

Uutrttt aUuufn/i'.su Tyler vV: Ansiis, 1^75 

i -VnunKlKlidiMO Dim ft*».' M*W I07(2t. I2'J M0_ 

lV f>WII s \I i |U7S) S|1lvil"-, uiolI|is uilhui llw 



\usirnlopapuan hyllcl frwg pawn t>itoJi(i 'Cscluidi \uu 

,/. A"*/., Slwpl, S<rn v ft3, 1-47, 

WAT>0Ki t., I ,. LlTTl I loitr,. \t J„ \\U«k J M .v 
RoUl.HrMjN, V. t \wi ) t onseiMilion status. eeoU^N and 
inaiiuijeinent ril the Spotted Tree lioj; | ftii'tia tye)U VhU 
It'ch. Hifli Stiit\ 116, Arthur livlah ht\t. Iin>n»i AVv. 
fippH Otf/A- EittlftW-i lleiik-lberu. Vttj M) i vi pp 



STRATIGRAPHY OF THE NEOPROTEROZOIC ARUHNA 
AND DEPOT SPRINGS SUBGROUPS, ADELAIDE 

GEOSYNCLINE 



By Ian A, Dyson* 



Summary 

Dyson, I. A. (1996) Stratigraphy of the Neoproterozoic Aruhna and Depot Springs 
subgroups, Adelaide Geosyncline. Trans. R. Soc. S. Aust. (1996), 120(3), 101-115, 29 
November, 1996. 

The Sandison Subgroup of the Lower Wilpena Group is unconformably overlain by 
the Wilcolo Sandstone and, together with the Bunyeroo Formation, comprises the 
Aruhna Subgroup. The Bunyeroo Formation is in turn unconformably overlain by the 
Wearing Dolomite which, together with the overlying Wonoka Formation, is assigned 
to the Depot Springs Subgroup. A number of subgroups in the Umberatana and 
Wilpena groups is also capped by dolostones that display similar characteristics to the 
Wearing Dolomite of the Depot Springs Subgroup. The dolostones are interpreted as 
having been deposited on major, sediment-starved hiatal surfaces under cold water 
conditions, each of which is adjacent to either a major incised valley or submarine 
canyon fill. The differentiation of these unconformity-bounded subgroups is based on 
their recognition as genetic units in terms of sequence stratigraphy. 
Key Words: Sequence stratigraphy, Neoproterozoic, Aruhna Subgroup, Depot Springs 
Subgroup, Bunyeroo Formation, Wearing Dolomite, Burr Well Member, Artipena 
Dolomite Member, Wilcolo Sandstone, Wonoka Formation, incised valleys, 
submarine canyons, dolostones, Adelaide Geosyncline, 



TStmaciimx qfjke Html SV«.tov ft f s - dftwt ' , ' )t)( ' 11 120i Ui WT- 1 13 

STRATIGRAPHY OF THE NEOPROTEROZOK ARUHNA AND DEPOT SPRINGS 
SUBGROUPS, ADELAIDE GEOS\ NCLJNE 

hy |A\ A. DYSON* 

Summitry 

Dyson. I, A. 1 10**6) Stratigraphy of the Neoproiero/oic Aruhna ami Depot Springs *-i»hpioups. Adelaide 
OnnytutmK Vnm ft tot S-AMi*\\W&k iflfo-), icn -1 15.20 No\emhei. io%. 

llic Sandison Subgroup uf the Lowet Wilpetia Group is uncouformably overlain by the Wilcolo Sandstone 
ninl. loitiMhei uiili iht- BuitvuMMi I'uriKuiHHi, ttiiiptiMj'- the Aruhna Sulnmwp. The BimytitiO Formation i> in turn 
hih 'nilurni;tbly tiverLttii by lilt- Wearing IJolnniilL- wlneh. In^etbcr with the ovei'hut^ WonoKu Formation, i* 
assigned io (hi' Depot Spinij.--- Subpfup. A luimhci tif >L»l\f r*mps in the I hnberafana ami Wilpcrw groups is also 
iappt-il hy dolosioucs that display similar characteristics to the Wearing Dolomite of ihe Depot Spring 
Snhyronp TllC dolostoues tire mteipn-led as having K-en deposited on major, sediment-starved hiatal surfaces 
tinder cold water eotnlit ions, etictt of which is adjacent to either a major incised valley tft flibtfiariiw canyon OIL 
PI.IC dillciVnltjliiMi it! these uncoiltonmly -houndeO subgroups w based olt 'tied PBCYfgnil tOft us yeuelte units m 
trnns ol scquemv slraligrapby 

Ki \ VYiams: Sequence shatiuuiphy, NeoptoLeiWoie Aruhna Suhgtoup. Depot Spring Slfogruup. Huuycron 
I -oi malum Wealing DQlUiUilU. Ihio Well Member Aitipena Dolomite Mcmhor. Wilcolo Sandstone. Wormku 
\ ormulion incised valleys, submarine canyon--, dolosiones. Adelaide (icosviuline. 



Introduction 

The sltaligruphic nometklaturc ol the Adelaide 
Gcosynclinc emphasises the distinction between 
chronostratigraphic and lit host rat (graphic units 
(Prctss IOM7a). The positions o\' (lie ehnmo- 
slraligruphtc units do nol always correspond to 
hthosiraiiginphic hounduttes. Some bthosirniigraphic 
boundaries are unconformities and therefore assume 

ehruflnstmtigraphic significance, while others arc 

niappable Ihhological changes v\' regional signiilcance 
iIYeiss IOK7a). These differences between the two 
stratigraphies can best be accommodated by adopting 
;i sequence straiigraphic scheme. It depends OH 
the recognition of nutppable rock units within a 
chroiiusl rati graphic frame work id repcliti\e. 
genetically-related strata bounded by unconformities 
or lltcir correlative eoulnrnuiies Thus, a revised 
strait graphic uomenelLiiure oT Neoproiero/oic 
succCsMoits in the Adelaide (ieosVOeline eotdd he 
based on diHerciitiation of subgroups within a 
sequence slrali^rapliie Iranievsvirk (D>son l l '^2a, b. 
I9%at. Forbes & I'reiss (I9S7) suggested dure 
v\iis merit in uniting related iteposttional units in a 
sinclc subgroup. 



• Nabtinal CtMIJTC Ibi IViiotLLiin Geututiy and Ueophyskv. 
Hiuveixily nrvXdetaKle Ailt.'l;tidc S Ausl. sOO.S 

' DvsitN, 1 A ( \W?S) Sedtmentolo^y and sduhpiaphy of 
the Neoproiero/oic SiiiutiNon Subgroup; a siorm-donu 
natrd slialluw marine sequence in the AticJilidc 
CirdvytitJine, Siinlli \tistra!ui PIlD thesis. Flinders 
t'liivrisilv nrSonih Aiisnalia liinpuh.)- 



hequence analysis ol the Umberalana (noup 
(L)vson 1002a, 1005 1 , 1006a, b) and Wilpena Gr»>u[) 
(son der Hotvh vi w/ I0XS: Dyson 1002b) has led l<> 
the rcco^nilion of several unconformity-bounded 
depositional sequences, hi a study of the Samlison 
Subgroup ( Dy^on I0OS 1 \ t stratiyraphic units 
immediately overlying tins sequence were examined 
in order to understand heller the spatial and temporal 
relationships of I lie Lower Wtlpcna Group. The 
s<indKoii Suhgroup is uneonunmahly *»veilain by the 
Wilcolo Sandslonc and togclhcr with the Bunycroo 
l-'ormalion is herein assigned to the Aruhna 
Subgroup. Similarly-, die Bimveroo Formation i^ 
uueonformably overlain by the Wearing Oolowilc 
and itvgeLher with the Wouoka Formation is assigned 
to the Depot Springs Subgroup. The Saridisoii. 
Aruhna and Depot Springs subgroups (Fig. I) are 
defined as geneUe umis thai ore considered major 
unconformity- bounded, depositional sequences in 
the sense of Milchum { 1077). Of purticnlar 
stguiliciuiee is the nature »>| the Wearing DolinttUe and 
its relationship to other NcoprtUero/oic dolosloues or 
units that contain appreciable dolomiic in the Adelaide 
Geosyncline, i.e., Nuecaleena Formation, lindelpina 
Shale. W'arcowie Dolomite and the Arlipcna Dolomite 
Member (new name) oi the Fnorama Shale. The 
n. niics ^Wilcolo Saiiilshme". "Aruhna Subgroup", 
"IXpol Springs Sli&g^Up^ and "Arlipena Dnloiujle 
Member" have been reset vetl by the Central Register 
of Australian Stratigraphic Names. 

Aruhnu Subgroup 

In the southern and cenh'ul Flinders Ranges, the 
ABC R;i,uge Quart/jte is overlain with lOOfil 



102 




LA. 


A M >N 










MORALANA 
SUPERGROUP 


HAWKER 
GROUP 

WILPENA 
GROUP 


POUh 




EARLY CAMBRIAN 


O 

M 
O 
ac 
uj 
i— 
O 
QC 
Q_ 
O 
UJ 


HEYSEN 
SUPERGROUP 

WARRINA 
SUPERGROUP 


ID SUBGROUP 


MARINOAN 


St 

< 
at 

< 

CJ 

< 

o 

UJ 


< 

UJ 

o 

<c 

_l 

UJ 

< 


lis 

S8f| 

l/J 


Wonoka 
Formation 


Wearing 
Dolomite 


Q_ 

< 3 

2 O 

x at 

3 o 
CK IB 

< ^3 

on 


Bunyeroo 
Formation 


Wilcolo 
Sandstone 


SANDISON SUBGROUP 


UMBERATANA 
GROUP 

BURRA 

GROUP 


AAAAAAAA 
AAAAAAAA 




STURTIAN 


TORRENSIAN 


CALLANNA 
GROUP 

'ALAEOPROTEROZC 


CUROIMURKA SUBGROUP 


WILLOURAN 


ARKAF 
)IC C 


I00LA SUBGROUP 




ARCHAEAN & F 


OMPLEXES 


Pre-ADELAIDEAN 



l\g 1 .Stratigraphy 6f the Auihna .Subgroup and DcpCgt .Springs Subgroup with respect to selected lilhosiratit>raprnc unils ot 
ihe Adelaide Geosvncline. Nuie ihe siraii^raphic position ol dolostones within the IJmberatana and Wilpena Groups. 



iI'im ont'ortnity by a thin {2-5 m>. massive. purple, 
coarse-grained to pebbly eioss-hedded sandstone of 

Fluvial origin (Plummpi' io?X). in places. H is 

intcibeddcd with conglomerate and purple shale. It 
is, in litrn. overlain by greyish ret\ shale and thin, 
inteiltcddcd lenticular sandsione of the Bunyeroo 
Formation with a sharp, conformable contact. Dyson 
(I9 ( i2b, 1995*) lecogmscd Ihe regional significance 
of this unconformity ami the nalure of Ihe channel 
fill taetes OVCflyiilg the Unconformity. The channel 
llll faeies is referred to herein as ihe Wilcolo 
Sandsione and is conformably overlain b\ shale »>f 
the Bunyeroo Formation. 

The Wilcolo Sandstone and Bunyeroo Formation 
together eonstitule the Aruhtui Subgroup (Fig. 2). U 
is a third-order eycla that is overall transgressive and 
was deposited during one euslatie tail and rise ot 
relalive sea level A reference section is designated in 
Bunyeroo Valley between A room) Ruins and Wilcolo 
Crock on PAKAC'IIII.NA. The Aruhua Subgroup 
was studied at Bunyeroo Gorge, Mount Terrible, 
Partacoona. Pcttana Gorge, Trebilcock Ciap and die 
Mouill fioddard and Angepeua Svnvhoes (Pig 3) A 
type section lot the Wilcolo Sandstone (Fig, 4 1 is 



designated in Wileolo Greek, 2 k|)i south of 
Bunveroo Gorge (lal. 31 2.V UJ" g, [i m g 13N |V 
I2"F.i. 



/.our/ Tttqiieuct! hotinifitn 

The Wilcolo Sandsione repic.--cnls an incised 
valley 1511 near Ihe lop ol I he ABC Range Quari/ile, 
A shallow palaeovaltev can be itaeed I'mm ihe 
A roan h Valley (30 in thick) to south uJ Buuvcioo 
Gorge where it attains a thickness of 3 m (Fit: 4). 
The base of die incised valley fill is interpreted h. in- 
a scqUOllCC boundary that was cul during a lowsland 
ol relalive sea level. Al Parlaeoona (Fig 3), ihe base 
of Ihe incised valley is interpreted as a combined 
sequence hoiindary/transgivssivc surface. A possible 
sec|uenee hounduty exists neat the lop ol die ABC 
Range Quarl/ile at Hidden Gorge iFig. 3). Here, the 
seqnenee boundary is overlain by a thick (> It! in). 
WSfJ coarse-grained sandsione or conglomerate thai 
is typically bimodal ami very well-sorted. Internally, 
diagenetic chert occurs as replacements and 
overgrowths. The same texture is observed in the 
Wilcolo Sandstone neai Bunyeroo Gorge. 



ARUIINA AND DEPOT SPRINGS SUBGROUPS 



HH 



O 

O 

INJ 

o 

GET 
LJ 
\— 
O 
Of 

LL 

o 

UJ 



MORALANA 
SUPERGROUP 



HEYSEN 

SUPERCROUP 



"V — v — v — V 



WARRINA 

SUPERGROUP 



-v v — "V V - 

ARCHAEAN 



HAWKER 
GROUP 



WILPENA 

GROUP 



UIYIBERATANA 
GROUP 



BURRA GROUP 



CALLANNA 
GROUP 



"V — v — v — v — v — V V V 
POUND SUBGROUP 



Wonoka Formation 



Wearing Dolomite 



Burr Well Mbr, 



Sunyeroo Formotion 



Wilcolo Sandstone 
ABC Range Quortzite 



Brachina Formation 



I~ Bayley Range Siltst. Mbr. 
I Uoonlloh Sitlsl- Mbr 



Mooiooloo Siltst. Mbr. 



Nuccaleena Formation 



Elotina Formation 



Trezona Formation 



Enorama Shale 



Artipena Dolomite Mbr. 



Etina Formation 



Tarcowie Siltstone 



r 



Cox Sandsterte Mbr. 



Hr-y y y y y y y 

Tapley Hill Formation 



j Wackt-rawirro Dolomite Mbr. 

Sunderland Mbr. 
iMt Caernarvon Creywacke Mbr 



Tindeipina Shale 



Wilyerpa Formation 



Warcowie Dolomite 

— y y rf y y V V— 

Holowilena Ironstone 



Pualco Tillite 
-v — v — v — v 



Skillogalee Dolomite 



CURDIMURKA SUBGROUP 



ARKAROOLA SUBGROUP 



to MES0PR0TER0Z0IC COMPLEXES 



EARLY CAMBRIAN 



MARINOAN 



STURTIAN 



TORRENSIAN 



WILLOURAN 



< 

UJ 
Q 

< 



< 



Pre-ADELAIDEAN 



Fig. 2. Unconformity-bounded subgroups conforming to depositions! sequences in the Umberatuna Group and lower 
Wilpena Group (after Dyson 1995' h 



I. A. DYSON 




KILOMETRES 



Fit;. 3. Tcclono- sedimentary provinces Of eastern South Australia, showing localities of stratigraphic sections in the Adelaide 
Geosyncline and their relation to other localities on the Smart Shell and in (he Torrens Hinge Zone (after Dyson 1995'). 



AKl UNA AND DHPOT SPRINGS SUBGROUPS 



m 



'''''■'■'■'■' 






I L 



500 



Wonoka Formation 
Wearing Dolomite 
Bunyeroo Formation 
Wilcolo Sandstone 
ABC Range Quartzite 
Brachina Formation 
Nuccaleena Formation 
Elatina Formation 
Trezona Formation 



E5SS3 



Y7777A 

p. "". ■* "t — S - ^" 1 

Y* * * * * *\ 



METRES 



Strike and dip 

Geological boundary 
inferred 

Type section 



47 



Fig. 4. Geological mapoflbc lower Wilprn:i Group ;it Runvcroo Valley si towing the lype set lion lorlhe WilcoloSaiuiMone 
(iilU-i Dyson l'WV>- 



lOo 



I A DYSON 




Fig. 5. Pebbly cross-bedded sandstone (2 m thick) of fluvial 
origill, assigned to the Wileolo Sandstone. overlying 
shallow marine sediments of the ABC Range Quartyile 
about 2 km south of Bunycroo Gorge. The* channelised 
b;ise of the sandstone is immediately left of ihe native 
pine in Ihe centre foreground. 



3 
O 

O 
CD 



I 

< 



0_ 

O 
a; 
o 

00 

=> 

CO 



o 

CO 

Q 
21 
< 

in 



BUNYER00 
FORMATION 



WILCOLO 
SANDSTONE 



ABC RANGE 
QUARTZITE 



2m 




outer shelf 

Ts shoreface 
fluvial 



SB 



^6 



grain size 

rill 



SB 

TS 



tidal flat 



inner shelf 

^ SCS 
^Q HCS 
^<^\ current ripples 
\y trough cross-bedding 
< < < herringbone cross-bedding 
\^kj wove ripples 
~~^^ lenticular bedding 
= horizontal planar lam. 
* * * clasts 

sequence boundary 
transgressive surface 



Pig. 6. Strati^raphic log of the Wileolo Sandstone at the proposed type section, south of Bunycroo Gorge. 



ARlJllNA \N0 DFPOTSPklNOSSl IJCROl PS 



K) 




Fig- 7. Fluvial channel :n ihe h:tse riflhc WUeoIo Sandstone 
iv overlain by .1 20 cm-thick mnlurc sand>lOnc thai is in 
turn gradnhoiully overlain h> shak of the Bwnyenjo 
hontuuion- The head o!' line hammer murk* the sharp coiv 
laci beiwecn the iwu sandstones, interpreted as the trnns- 
pressiw surlact- 



consists of interhedded metre-thick, pebble \u cobble 
conglomerate, medium to coarse-grained sandstone 
and shale. The conglomerate and sundsionc display 
planur -rabuiur cross-bedding and SGS respectively, 
and are interpreted as having been deposited in u 
shorefnec euvironineiu within an incised valley Mil. 
About I 5 km enher side of Tiebileoek Gap. the 
incised valley till contains large (500 x 100 mi rafts 
of drapine breccia. thought l«» have slumped inio the 
incised valley dining the early stages ot deposition. 
On the south limb ot the Mount Goddard Syncltne. a 
greyish red T fine-grained santlstonc crosivels overlies 
Ihe I'lupa Siltstone. The sandstone, about 1 m thick, 
contains angulnr to sub-rounded clasts ol diapiric 
material suggesting exposure of a nearby diapir and 
is interpreted as being ot lluvial origin (Tig- S)_ 

At Pettana Gorge (Fig. 3), the W'ilcolo Sandstone is 
absent but Tor a thin remnant of gntl> and gossanous 
sandstone It is erosively overlain by a boulder 
conglomerate at the base o\' a submarine canyon in 
the Wonosa Formation I Dyson 1995*). ticil mapped 
previously on ORROROO (Binks I9oS). The base of 
the Wdeolo Sandstone is not exposed at I lallett Gove 
(lig- 3U but at Mount Terrible il is overlain by 
interhedded greyish red siltstone and mature 
sandstone At this locality, the base ot the W'ilcolo 
Sandstone is interpreted as a combined sequence 
boundary/transgressive surface, A similar situation 
exists at Finke Springs on the north limb i»t the 
An^epena Synchne where a thick-bedded, medium- 
grained, swaley cross-stratified sandstone ol' 
shore face, origin overlies the ABG kange Quarizite 
On the south limb o! the Angepena Syneline near 
Sliepherds Bore iFig 3). a dccimetre-thicL lidally 



Wihuitt) Sandstone 

"flic lluvial channel at Ihe base i)i the Wilcolo 
Sandstone (tag. 5) near Bunycroo Gorge is overlain 
by a thin <e. <l mk mature sandstone (Figs 6. 1) that 
often displays swuley cross-stratification iSCS). 
hummocky cross-si ratification (IICS) and sym- 
metrical ripples (Dyson 1992b). The base of the 
swaley cross-stratified sandstone is mierpretcd as a 
li'ansgrcsMve surface. At Partacoona. an un- 
conformity at the top of the ABC Range Uuart/ile is 
overlain by 25 m of mature, off-while quart/itc that 
displays trough cross-bedding of tidal origin and 
large symmetrical wave npples with abundant well- 
munded clasts of gravel to pebble size. The ciuarl/Jte 
was deposited in a possible incised valley of similar 
dimensions to that observed in the Aroona <i\\<\ 
Bunyeroo valleys. A contact with overlying sbale of 
ihe Bunycroo Formal ion was not observed- Near 
I rcbilcoek Gap west of Be It ana. Ihe Wilcolo 
Sandstone varies in thickness from 20 50 m wheu it 




Fig. 8. Conglomerate fmrii ihe Wilcolo Sandstone on the 
southern limb of the Mount Goddard S\nchne ti con- 
tains carbonate clasts of possible diapirk origin Coin is 
2H Mini in dianieier 



I OK 



f A PVSON 



ctoss-hcdded saudslone erosively overlies, the Ulupu 
Siltstone. ll is overlain by a M) m-lhiek section oi 
iMicrhedded greyish red shale and sandstone lhal 
vrades upward into reddish shale Of We Bunyeroo 



The Bonycioo Formation (Dalgarno & Johnson 
1964 1 ^ 700 m Ihick in its type section ,(( Br.icluiu 
Gorge (Fig. Jf. when- il consists nf laminalciJ IrJ 
mtjSSlvtf. dark reddish brown Ti;ile_ The i>V< i I 
U|«M itut-fiiiiiU' succession is punciu.Hcd fcj .1 scries ol" 
subtle, upwatd- coarsening, cycles lhal in places nmge 
I10111 5-10 m llnck. Sedimentary stuicturcs 
associated with very fine to Juic-erainc'd kancfiftffflc Ul 
Hie lag ol same" cycles include small scale pfl ■■■. 
hedding and micro 1ICS. 

The Bunveioo l-ormation was tor I he mosl pan 
deposited below slonn wave base Ml a middle 10 
outer shelf selling, Dyson [l,«W2hj placer! the 
Huuycmo Formuiiou in a Uanstjre-ssivc systems nacl 
thai was capped by ihe former Wealing Dolonnlc 
Memlvr of the Wonoka Formation, 



The sequence boundary at die top of the Bunyeroo 
loiiiuiljoii is eoitictdenl with Hie former Wearing: 
Dolomite Vkutbcr ofThomson (l%5». It is elevated 
herein to formation status to reflect its regional 
.significance, Deposition ol' (he Wearing Dolomite is 
interpreted &$ having been contemporaneous with (he 
canyon unconformity til ttie hase of the Wonoka 
lormiition (Dyson KjSS". 1906a. hi. 

Dt fmSttfttnttl nnitn/iltirtll 

The Wilcolo Sandstone was deposited in a fluvial 
anil csluariiie to shallow marine environment. The 
Bunyeroo formation was deposited in progressively 
deeper walet in J middle to outer shclJ' selling aiul 
constitutes a transgressive systems tniei. Thus. 
sedimentation ol the Aruhna Subgroup was unable to 
keep up with subsidence, resulting in a tleposilional 
transgression m the sense ol Curray tl%4), The 
Bunyeroo Formation thickens eastward ol the 
Torrens Hinge Zone {Fig. 3). Adjacent to diapirs, 
onlapping sediments of the Aruhna Subgroup arc- 
thin, ilowevci. localised (hick development of the 



Musis, P.W- il')K7) r.uhonnte shell ind h.is.n 
srdiuitnhiiinn, late Protcro/oic Wonoka Phonal him 
Souih Australia. PhD thesi.i. tJnhetsav >»t -Vlrlaide 
lunpulvi. 

1 11 KosA. P. \. 1 .198**1 Gl-uIoc-il hisloiy M.s|UCnce 
Mralutinphy ol ilk- lute Proiem/mc Wonoka formation. 
MtHltn-oi VlnnK is tt.irij.es South Aiislralw. Phi J Ihests, 
I linden Fnivcr.ilv ol South Australia oiiipult ), 



Bunyeroo Formation occurs adjacent lo some 
diapirs. A thick, black succession of sulphide-rich 
shale adjacent to the Mucaloona Diupir (Coats 1973) 
suggests anoxic, deep water deposition of ihe 
Bunyeroo Formation, perhaps associated with die 
formal ion of a civstal utabeu over the diapu due to 
sail depletion. The diapus of the Flinders Ranges 
"Men contain volcanic \enodasis (Pteiss IV87h). ami 
Cnnl£ (1973) suggested that many diapirs on 
GOHjRY were active aiulcKposed during deposition 
of ihe basal Bunyeroo Formation. An inferred 
voJeanic componeul of the redheds (Muwson 1939 
Plununer l97Sa) may be iclated to depositkmul 
Oilinp of Bunyeroo sediments adjacent to i^-pri$i>J 
duipiis. 

Dahymple [liW2] suggested lhal cstuarim' 
sandstones weic transgressive in origin because 
estuaries owed dieii existence lo marine [laiidlllfi ol 
incised valleys. On the oihcr hand, txxon researchers 
icy,. Van Wagoner ,7 ui I0K7) aioiied Ilia! Iluvial 
sediments at the base ol" incised valleys should K* 
assigned lo (he lowstand systems tract dtfposliod 
during an initial fijll and subsequent early rise of 
relative sea level. Alternatively, sueli Iluvial llflr(S 
may be the updip eo;uivalenl of tiausyressivc tn;n uic 
sandstones. The lack of beach deposits between the 
fhnial and esluartne sandstones of the Wileoto 
Sandstone at Bunyeroo (iorgc suggests Hull die b..se 
ot iltc esluuiinc sandstone is the transgtessjve 
surlaee Mil ensuing transyressiou eroded and 
.vwoiked the former beach sedtmenls. 

Depot Springs Subgroup 

The Wonoka Pormalion <Dalgarno Ul Johnson 
Iv»b4) and Wearing Dolomite together represent a 
lransgressive-reyressi\e (TR) cycle lhal is reletm! 
to as the Depot Springs Subgroup (Fig- I ) The f Vjiot 
Springs Subgroup eonMitUtes an uncontornulv- 
boyiWg deoositional seijuencc and was studied ai 
a numl'o ol localities 0,1 PARACHII NA, COPLFY 
and MARREE nieluding Peltana Gorge. "Wyaeea 
Blull. Brachtna Gorge, Bunyeroo fiorpe Beltana 
Ilill Mount Cioddnrd. Shepherds Bore and 
Mundaweriitia Well (Fig. 3j_ A reference scelii»n for 
the suhgroup is designated in and adjacent lo 
the Palsy Springs canyon ol the Aneepena Syncline 
near the Depot Springs IIS, 40 r.m east ol 
Copley (Fig. it 

The base of the Wonoka Formation was mapped » -n 
PARACHII NA (Dalyumo & Johnson 1966) wher. .. 
cohuu change occurred above greyish red sandstones 
at the lop of the underlying Bunyeroo Formation, 
This boundary corresponded to a rather abrupt 
increase in lime content. Gostin & Jenkins (I9K.H 
del med a dcehnclre-lhicK doloslone overly inp 
reddish shales ol the Bunveroo Fonrmtion. refened 




ARl 'UNA AND DEPOT SPRINGS St KGROl 'KS 



LOU 



pig. l ). Sharp Contort between the Bunyeroo Formation and 
Wearing Dolomite ill the Angcpcini Syncline. The 
Wearing Dolomite is aboitf M) em thick. Us ha&e is 
marked h> the head ol the hammer with sedimentary lac 
iti£ to the right. 




Fig. 10. Wearing Dolomite displaying rnicro-HCS, 

Angepena Synchne. 



lo tnfornuilly as the Wearing Dolomite MemU-i 
(Thomson 1965), as the base oT the Wonoka 
Formation. Jenkins (1993) defined the base Q$ the 
Wonoka Formation in Bunyeroo Gorge at the base of 
an intnifin irrational conglomerate within the Wearing 
Dolomite Member Haines (I9S7 : » divided the 
Wonoka into 1 1 litholneics units. These units were 
subsequently adopted by other workers (e.g.. Di 
Bona 1989*: Chrislie-Blick trt til. 1900) with the 
prefix \V. 

fj'ut'r sequBtttt boundary 

The base of the Wearing Dolomite is tlellued as a 
sequence boundary. Il can hi* cither sharp or diffuse 
in nature. Jenkins (1993) interpreted a sequence 
boundary at the base of an intralormational 
* i n-'lomcrale within the Wearing Dolomite. 
I iowever, the iniiaformational conglomerate displays 
edgewise clasls thai have in ihe past been interpreted 
as storm rpSCttes (eg.. Dyson &. von dct Botch 
19X6). Dyson (1992b) suggested that the Wearing 
Dolomite represented deposition within a condensed 
section that included a possible maximum flooding 
surface, Furthermore, the Wearing Dolomite vvas 
deposited on a sediment-starved hiatal surface below 
storm wave base (sec below). 

Wearing Dolt unite 

The former Wearing Dolomite Member ol 
Thomson (1965) is a thin cream, dolostone or 
dolomitie sillstone that has been mapped ovei 
extensive areas o\' the Flinders Ranges (Forbes & 
Preiss I9S7). II corresponds to unit I of Haines 
{ J9S7-), The Wearing Dolomite often sharply 
overlies the Bunyeroo Formation with apparent 



s*£ZZ?£2^ 




'.':' : I.- 




WONOKA 
FORMATION 



WEARING 
DOLOMITE 
Unit W1 



>**gm 



UnM VVJ 

Ihln bedded tempastite tacias 

Unit m 

'wall plaster' do'ostone tocies 

QPL A HCS ternpeslitB IocIbb 
Burr Well Member 

stromo!oll"t llrriBslune iaoies 
SCS sands'one facies 
Cap Dolnstonu Member 

laminaled dolostone loctea 
Canyon fill Memoer 
'wall plaster' doluslane fades 

slump laclfls 

cross-bedded pebbly sandstone facies 



l-'ig- 1 1 Schenuilte cross -section through the Patsy Springs Canyon m the Anuepeila Syncime (after D\son P095 1 1. NOW 
the relationship beiwecn dpi ok tones t>l the Wearing Dolomite. 



I Ml 



i \ KftSON 




% 



I i^ I.!. Hrcc t uOa luncsiohe nP ihe WtMnnp Dolomite 
uncoiilonn;ihly overlying ihc Bimvcroo formation rjisi 
ul IklUtna I hll 

conlormiiy (fig, 9). It is characterised by wavy «J 
parallel lamination and, less commonly, by micro 
HCSffig. 10r A similar situation exists on the south 
Itllflb Of the Augepena SynC line and north limb of the 
Mount Goddard Syneline. Near Mundawertina Well 
and on the northern limb of die Angepena Syneline, 
(he Wearing Dolomite splits into two thin dolostoues 
thai aiv separated by up [Q 2? m ol taminaied grey 
greed --hale. Al Pellana Gorge (Fir:- ^). a 2 m-ihn k 
boulder conglomerate cuts downward into the ABC 
Range QHiail/.ite. Lt marks the base of a shallow 
canyon with an estimated relief ot 7" m About 5 km 
souih of l J elhuiu Gorge near Wyacea Blul'l". the 
Buuyeroo formation attains a thickness of about 50 
in where ft is sharply overlain by a decimetre Thick 
dolostone of ilie Wearing Dolomite wiih a sharp 
planai coniael II. is interpreted as the correlative 
conformity adjacent to the submarine canyon, 

Other dolostones are also genetically rclaicd to the 
Weaiiug Dolomite al this slraligraphie level (fn;. 
1 1). About 2 km casi ol Bellana Hill ff'ig 3| ( ihr 
Wearing Dolomiie passes laterally mm a several 
uictie-ihiek. cream to orange laminated dolostone 
that i% often nroccialed and present as detached 
slump blocks iLecson ,V Nixon |9oo>, Here, it 
overlies reddish shales of the Buuyeroo formation 
vi ih -m augnlni unconformity (Pig. 12). This 
dnlnstnne is thought to represent the "wall plaster 
veneer" ot lackoil ( I9KKM A similar relationship 
can also be observed near Muudavverliua Well, and 
also on the south Innh of ihe Angepena Syneline 
tvhCft the Wearing Dolomite can be traced into 
slumped wall plaster at the edge ot ihe Patsy Springs 



lilCklU'ii. KB iOKHi ricol.mi.^il ||iKi.>ry , t n.j rtffglll nfllM title 

f'i"ii'M/ini' fttriirt" Hiu Gfonyan QirBpWx. AdctMiik 

<«>i>%yivlin.., S, \. PWD Itvbin, Mimlcrs I nivrf-iiv <-i Smith 

Au-.ii.iiin iirnnuha, 



canyon iDyson 1995'). On the soiuh limh ol the 
Mount Goddard Syneline. ilk- Burr Well Member ol' 
Di Bona (I9N9'J consists of mature, medium 
grained, swaley cross-stratified, carbonate-cemented 
sandstone or iniraformalional conglotneraie with 
edgewise elasls of dolostone. and passes laterally 
uiio stromatolitic dolostone (see Kg. 1 1 1, A basal la; T 
comprises duipiric detritus. Where the Wearing 
Dolomite displays such unconformity, it is defined as 
the Burr Well Member in the %cnse of Di Bon,. 
(19X9 i 

The base of the Wearing Dolomite is interpreted lo 
he j. deep watei sequence boundary, A maximum 
Hooding suilacc al the lop rjf the Buuyeroo 
formation may coincide with (his sequence 
boundary. The ditTusc bas L ol (be Wearing Dolomite 
suggests thai early post-deposttional doioniiti/alion 
took place on a sediment-starved hiatal surface. 
Wavy lo paraltel lamination and micro- DCS further 
suggests deposition below storm wave base (e.g.. 
Dyson 1995). The Wearing Dolomite can be traced 
into the Burr Well Member on Ihe south limb ol Ihe 
Mount Goddard Syneline- The Burr Well Member 
was deposited in a storm-dominated shnrcface lo 
tide-dominated, lagoonal environment Its SrUlTp. 
erosional base is a enmbined sequence boundary/ 
iransgressive surlace. The "wall plaster veneer" 
was deposited on the canyon shoulders. It is 
interpreted to he coeval with deposition ol llv 
Wearing Dolomite and unit W2 ot* the Wonoka 
Formation (He. 1 I ). 

The Wearing Dolomite, logelhcr with the Wonoka 
formation, is present on the Stuart Shelf where it c;in 
he observed in drilleoTc (&£., Bopeeehcc 2 al 3o72 
mi. A possible Wearing equivalent, only a lew 
ceniimctres thick, crops out south of Bills Lookout 
near ihe north- -western >ule of Lake Torreus within 
what Mas previously interpreted as Varloo ShaJe 
Uohns vi uL 1966k I his suggests lhat the Wearing 
Dolomite transgressed die Stuart Shelf prior (it 
deposition o| die Wonoka formation 



Wtmpko for/nation 

The Wearing Dolomiie is overlain by unit W2 ol 
ihe. Wonoka formation, consisting of greyish iv\\ 
Imc-giained sandstone and inlerhedded calcareous 
shale. Il grades upward into the dominanily 
calcareous shale of unil W3. 'Die succession 
represented by units W2 and W3 is overall 
tt'ansjjressive. A colour change to giceiush 
limestones in the middle section of unit W3 marks 
die base of regressive sedimentation in the WOnobil 
hormation I fig. 1 1 ). 

The hthofacics units W3-7 inclusive of the Wonoka 
foimalion display an overall upward sanding Ireml 
It culminates in a thick, stornvdommaied mixed 



AUUHNA AND DH'Ol 

UH'bonale/silieiclaslic succession, commonly displaying 
QtTS t Haines l°W) ilua was deposited in an inner K1 
middle shell selling. Units WW ot the Wonoka 
l : ormaltoii. as interpreted h\ Haines and oilier 
workers, represent regressive sediinetiuuion from 
iniliiil deposition he low slonn wave base lo 
deposition above fairWcalfiei wave base. 

A nnmber of nurlre-lhick. greyish red. medium - 
grained sandstone beds in tinil VV7 dial ufs 
characterised by SCS may represent forced 
regressive deposits associated with a falling sUl r' c " 
systems naet (£.£., Hyson IWfv>. In the Anucpcna 
Syneline. a discontinuity at this sltaliexaphit level \$ 
o\erlain by a several metre -thick medium-grained 
sandstone thai in many places displays hori/onial- 
planar lammalinn :.<ul S( S_ It is interpreted to 
represent deposition on the lower shoelace in a 
murine environment- Pi Bona ^ von der fiorch 
i IWt inletprcicd a low Maud delta a( this level in (he 
I iinheraiana Nvncliuc\ I'nil VV7 is oscilatn by a 
succession ol shallow uiaiine sandstone and 
< -arlmnaic ol (idnl and lugoonal origin (Haines 1990) 
that coiresponds lo units WX- 1 I. < Occasionally, Htae 
units can be obscivcd lo yrade upward into the ted 
shale mm\ sandstone ol' I he Boitney Sandstone c .£., 
near Nbatut (Jnddard. 

The IK cycle ol the Depot Springs Subgroup is an 
unconformity-hounded, third airder tlcposilitui.il 
sequence. The canyon llll. represented by units Wl. 
v\\' himI io a lev-,ei extent W3. constitute the 
tians;jicssi\c <vcle ot the Upol Npiines Suhvrouiv 
IVposaiou ol unit VVI in the canyon bit w .1.-. 
contemporaneous with deposition ol Hie Wearing 
Dwloiiulc (Hi a sediment-starved hnttal sulfate A 
,m,, ihlc condensed .section is represented by unit 
\V 1,Tln inoUlk ol this tmii corresponds to an abrupt 
colour change and increase in lime content A 
maximum flooding surface nm\ be contained within 
uml Wt and is theieloie equated with a dmvnlap 
siiifneo ll is ovi-rlain by the recessive hthnfaeies ol 
Hie Woimka |-oiiuali<>n. 



Sulfiiuimii' untsaus 

The Wearing Ooloinite was developed adjacent In 
a major submarine unconformity on a sediment 
Mai ved hiatal surface that corresponds tO •' combined 
sequence boundary and major floCKllttg surface 
(Dyson IWS 1 .), It can be obseived to pass laterally 
into slump breteioliis on (he shoutders of submarine 
canyons., c.y.. neat iicllaua Hill south limb of 
Ancepena Syuclinc. On ihc north limb <>l the 
Anecpi-na Syneline. the Wcarine Dolomite consists 
ol Iwotlotosloucs that arc -.cpaiaird by some 20 m of 
shaltv The upper doloslone overlies the truncated 
>T M r ill a lower dolosloue. sueecstine cauym 
erosion octttrred before deposition o\' the nppei 



SPKIMiNSl BGKfil in HI 

doloslone- Furthermore, tinil \VJ overlies the fill OJ 
the Palsy Springs eanvoii til the Aniiepena Syiiclnic. 
Retrogressive slumping on die ottlei shell' was the 
precursor lo canyon incision. and proceeded up lo the 
le\el of unit \V3 in Ihc VVonoka fV»rmati"U winch 
marks the turn around from Iranstiiessive lo 
regressive sedimentation in the Wonoka bormntion 
(Ha 1 1 1, This interpretation questions the inning of 
earliest canyon incision expounded by other workers 
thai coincided with deposition of units W3. W4 01 
\Vs ( c .£> Haines I9M7- 1 ; Di Bona |Wft Clinslie- 
Mukentf, \mi fcBMl. 



I /<!>,' r wtfuvm r htHHhliirv 

The fjoaney Saudsume often dispta\s a sharp 
though apparently conlorntablc contact with the 
under! vine Wouokn I 01 malum, Hovwu-r. -he 
relationship becomes disconforrnablc tn the vicinilv 
tJl dtapirs. Adjacent to some diapiis. c»|L$ Rnrirt 
Diupir 4Mi (/OI'Lh'V and I'tome Diapir on 
PARAnill.NA. (he Homiey Sandstone dt-pl'fVs .m 
uncon I (finable relationship with the WbTiuka 
formaiion (Dyson unpub ). Thus. Ihe base of the 
Btmncy Sandstone of the I'ouint Subgroup is 
iulcrprelcti us a sequence boundarv. l>_\son 1 I'W 
iQVfifl. urii'iiib 1 siuj^esied lh.it the development o! 
severiil prominent uueonh Mini ties \>r sctpicnce 
boiindaiies wilhm die Umberatana anJ Wilpcn.i 
fioaips was asscvialed with periods ot iuii\e and 
passive dupuism. which in turn was related to major 
cUciiMouai events duriue bnMkaip of id. 
Neoproloi-o/oii sni>eicoiitiucul 

IHsctission 

Tin Ltttfq/ s^'C""/"^"" W{\ "itffivittv ht'iifhlni tmii 
The previous use of mi be roup m a Inlm 
•aiutiLiapluL sense has caused problems m regional 
mappiui: programmes. A £<>od example is Ihe 
di lercntialion ol inierplacial deposits in die 
I'ntberatana liroup, They etnhr.ice ihc barina 
Subgroup (Coal* vi uf. WW); Thompson |y6 c » m& 

the Willochra Suberoup tllio,,. «mi |.^0j, \ Uc 
intenlion ol the term farina Sub^mup wa-- n> nicl"-k 
all relitli\'cly deeper water sediments, in 
contradistinction from the domuumlly shallow water 
redbeds of the Willochra Suberoup (Coats & Pretss 
l L iS7j, 1 here has been inconsistency in application o! 
the terms, especially m some tunsitiona! regions 
where facies iiitertoiiguc or are uilerc;i]aled (dnils & 
I'reiss l l J87f Ihesoulhem portion olPARACHII N\ 
is such a region where a l>asis lor this disrinction is 
wan auied. I he re fore, a revised straliytuphic 
noinenelaiure of (he inierglaeial deposits could be 
based on dillcicnnahon .»t .iiheioui--. as a eenciic 
mill lit tei us ot scttueuce stiati,-;i^i:l" 



112 



I A 



A sequence straUgrapliic framework fur lite 
Umbciataiia and Wilpena groups is shown in Fig. 2 
unci is bused on differential ion of" subgroup* us a 
genetic tutir. An uncoiitormuy-boLiuded sequence 
incorporates (he Tindclpina Shale and TapJcv Hill 
Formation Similaily. an imcoulormil.v -hounded 
sequence is defined by Ihe lop ol the Tapley Ifill 
formation lo near the top oil he Ftina Formation. and 
an overlying sequence is eapped by an unconformity 
al die base of (he Hkuina Formation, The Willoehru 
SubgVOlIp pieviously included Ihe Marmoan glacials 
ol ihe Glut in a Formation (Thomson I959J. The 
I'laHiia Formation is euppetl by dolostone ol' the 
Nuccalcena Formation, die basal unit ol the 
Snndison Sllb£i*OUp. Ihe Sandison Subgroup is 
unconformnbly overlain by the Aruhna Subgroup 
lhai comprises the Wilcolo Sandstone and Bunycioo 
Formation. The Aruhna Subgroup is in turn 
niKonfoimaMy overlain by Ihe Wearing Dolonuie al 
Ihe base ol Ihe IVpot Springs Subgroup (Fig. 2). A 
sinular scheme was proposed tor the Kanmanioo 
Group of Cam&rian age (Dyson IW). More work 
is required lo differentiate genetic units within Ihe 
Pound Subgroup vvhieh contains possibly two 
■ ■in on I'M nut v -bounded sequences, wilh sequence 
boundaries al the base of the Bouncy Sandstone and 
F.diacata Memhei ol the Kawnsley Quart/lie (e.g., 
Dyson. IW).- 

IJthostraligraphic, hioMiatigraphie and ehrono- 
-haiigiaplue uuils will eonlinue lo be used as die 
basis lor most slraligraphic studies. However, the use 
of unconformity-bounded units is invaluable m 
basins where ihe development of stratigraphie units 
was controlled by tectonic episodes and custalje sea 
level cycles. In sueh basins, e.g., the Adelaide 
Geosyncline, unconformities pass Intern I ly into 
correlative conformities where traditional strati- 
graphy is unlikely lo differentiate the lateral and 
vertical extent of genetic units ElftflVe and below Ihe 
sequence honudaiy. I 'litis, the use ol uuconlormily- 
hounded units can contribute lo the understanding of 
the stratigraphy and yeologte history o\ a basin, it 
can provide Ihe framework for regional slraligraphic 
framework and it can enable ihe mapping and 
expression or slraligraphic concepts ihr which other 
stratigraphie units are inadequate (Salvador I'Wl. 

A hierarchy of uneonfonnily-boundctl units can be 
humiliated by determining sequence order in a bosin. 
Milchuin &i Van Wagoner ( 1991) proposed a 
sequence boundary hierarchy of live orders on the 
frequency Of boundary occurrence. Alternatively. 
Emhry (|Q93) suggested five orders ^ sequence 
boundaries hased oil the uatuie of die boundary. The 
latter method is possibly less subjective. However, if 
every pair of unconformities is used to recognise and 
name an unconformity-bounded unit such as in the 
case of ihe Cardium Formation in ihe Cretaceous 



DYSON 

Western Interior Sciiwuy ol Canada (Walker 1 *><><>>. 

Ihe stratigraphie uriils would grow unmaiiageahK 
(Salvador 1994). The use of die subgroup as a 
deposit ional sequence of third-order ey el icily 
thereby usefully limits the establishment id" 
meaningful and useful slraligraphic units on a 
regional and intei- regional basis. 

Tht \ixntftc<tinc <>! dolmtoncs capping Atfelnidium 
M'tfiicnccs 

The Wearing Dolonuie of the Depoi Spring* 
Subgroup was developed adjacent lo a major 
submarine unconformity on a sediment -starved hiala! 
surlaee that corresponds lo a combined sequence 
boundary and major flooding surface. There arc other 
dulosioucs or units eonlaining significant amounts of 
doloslone dial are associated wilh sequence 
boundaries ill the L'mbcralana and Wilpcnu Groups 
(Fig 2) They are the Watcosnc Dolomite, the 



5 



- 



a 



*c 



5 - 



10 - 



-15 





i 
4 

A 
Warcowie 
Dolomite 

3 Tindelpina 
A Shale 




SAMPLE LOCALITIES 




1: MUNDAWERTINA WELL 
2: -SHEPHERDS BORE 
3: UPALINNA DIAPIR 
4: WILLfPPA ANTICLINE 
5: ARTIPENA HUT 


5 


2 

J 

We 


[ 
Nucca 

L . r i •» !_orrT1 

anna Dolomite 

▲ i 

1 Sil 


A 

Artipena 

)olomite Mbr. 

teena 1 
3tion A 

lupa 1 
tshone A 



-10 



<5 13 C PDB % Q 



Fig. I I Isolopic rffttfl foi Adclnideuii dnltrslones (blWOtl roi 

Dyson 19050 



AIUUINA AM) DI-.POT SPRINl iS SIMKjROIjPS 



J 13 



Tindelpina Shale and the Muce.alcena formation, all 
of which cap sequences, til' glacigenic origin. They 
arc commonly referred lo as cap dolosloncs te.y.. 
Williams 19QQY Another dolosione occurs wilhin (be- 
lower Hnorama Shale anil ispronnnenl i»n souih-east 
PARAC.HILNA. c g., between llie Willippn Anticline 
and Marlins Well Dome- ll is formally referred III 
here as the Arlipena Dolomite Member of llic 
knotuma Shale and a type section is designated 2 kin 
south of Arlipena Hut (Kg, 3). Here, n consists rji 
two thin < 10 5(1 cm) doloslones thai are separated by 
* m ol greyish red shale, and mark- (he transition 
from tra regressive to overlying regtcssi\c 
seduneiiiauon ot the I norama Shale. 

These dolosloncs of the Umberalaun and Wilpcna 
•jumps display similar characteristics* and are 
interpreted to ha\e been deposited on major Hooding 
Miilaivs under -cold water conditions. Their isotopic 
data, based on Dyson ( IffflS'l in Pig. ' J- display two 
apparent (tends. The carbon i%oiopc values show a 
shift in nunc negative values with possible inctva-e 
m walei deplh. accompanied by an increase in 
rliagenesis This suggests that the Wearing Dolomilc 
was deposited m greater water depths than the pjhjjf 
dolosloncs and was most susceptible la secondary, 
potfl ileposiiioiial alteration, the oxygen isotoi e 
values show a ship lo more negative &'■'() ttbC'VO the 
siralieraphic level of the W'arcowic DoloiTnle. U"5 J CJ 
is scusinv'c to temperature changes, then il might 
show a similar trend Lo relative water de| in as 
suggested by o 1 '■(.' (e.g.. Baum tf al. 1994). However, 
the shifl lo iiioiv negative 0"O above the V^fWWK 
Dolomite is interpreted to reflect the overall increase 
mi palaeoteniperatiire following the Stnrtian 
glaciulion. This interpreiatiou must be viewed Willi 
cauiiom particularly with respect lo the Wearing 
Dolomite, because of oveipiiiils associated with 
secondary, posl-dcposiiional alleration. Deposition 
llf the Warcowie Dolomite in a glacigenic 
environment is suggested by die prcsenrc ot 
dropsU.ne-* (Dyson 1993', 1996b). Palaco- 
tcuipcralurcs lor the Neopiolei'o/on. dolostones arc 
thought to range from 5 C POT the Nuccaleeua 
Formation ami Wearing Dolomite, lo -5 1 C lor Ihe 
Warcowie Dolomite (C I* Kao pers. comm, 1995), 

Kadi ill" the doloslones was deposited on a 
maximum flooding surface associated with 
terrigenous starvation. "Ihe Mdendetla Limestone, a 
carbonate ot" Cambrian age, occupies a similar 
siratiyraphic position in Ihe Kanmanloo Group 
Incised valleys arc associated wUh the Milendelia 
I uncslone and SeachlV Sandstone (Dyson & von der 



Boah 1994: D\son 1996.1)- Progradiug slope 
complexes are assoeiaicd with the Tiudclpmu Shalc. 
Sueh unit*- arc considered lo be ihe downslopc 
equivalents of incised valleys iMilclium d it! |9 U 5». 
A pertinent question is why dolosloncs nl ihe 
Wearing Dolomite and Warcowie Dolomite do not 
appear to be associated with incised valleys that 
show donmumtly shallow water features. Major 
eMensional events coinciding wilh deposition ol 
these units, together with high rales of subsidence, 
resulted in no fall m relative sea level and piecluded 
development ol saudv highstand factcs. Instead, 
dolosloncs cap relatively d^\> walci sediments dtf the 
Bunycroo Formation ami Holowilcna Ironstone 
lespectivcly. Each dolostone is developed on an 
hiatal surface Lhat represented a period ol'lerrigenous 
starvaiioii. In each case, ibis surface can lv followed 
itilo Ihe submarine uncoiirormily where a major 
canyon was possibly cut on the outer shell- lit such a 
scitiug. a fle.unul response ot the continental margin 
may have occurred as an isosiaiic readjustment ip 
canyon erosion (Medians , r ,//, lovsj. Tor a wide 
continenlal shell. flooaral uplift ol the outer shell 
would nol have influenced ihe position of the 
shoreline, resulting in an erosional unconformity 
developed only across llie diMal shell (McGinnis ct 
«} l°'J3) The hiatus u.enerateil Wuiild be greatest 
across the Uiseul shell and decrease m a landward 
direction. Incised valley Fills ol the Seaclill 
Sandstone and Milendelia l-tmcsioiie are overall 
tiunsgressivc but display relatively shallow water 
IWaures at their base. Sequence boundaries at the 
base of Ihese mused sallcys were formed dUl'iflg 
lesser eMensional cvcnls when Ihe degree \>\ 
subsidence was relaiively small or. more likely, when 
the width of the palaeoshelf was relatively nanow. 
lhiK_ a tall in relative sea level on the outer shell 
resulted in subaerial incised vallcvs comprising I'.r.al 
fluvial lo esluarine tlcfjosils. 



Acknowledgments 

This paper is based on lield studies uudeitafii' 
during p<istgraduale research aL blinders l.ini\crsily 
under the supervision ol Prolessor Chris von der 
Rurcb. Several useful comments were suggested by 
revieweisrhris Nedm and Wolfgang Preiss. Aspects 
ol the currenl study were discussed with Peter Reid 
and Wolfgang Piciss. Oail Jackson and (iha/i 
Kiaishnan kindly dialled die figures. 



1 1 



I A HYSON 

KtTervntt's 



[t\uM ( ,!, S.. Uai'M. U, R.. Thompson. R R. Si HUMPHRY 
II). it 1 )').!) Stahle isotoptc evidence lor relative and 
etiMatic .mm level chanpes In tuiecue lo miynceue 
raibonaicv Baldwin C'oiiiUv. Alabama. (><W. \ f >,\ dfflM 
Hull. 10<>. s:i-S.v). 

UiNKs. i' I. I, I'lfiKj QfcRQRGO ftifcip sheet. Geological Alius 
n| South Ausiiaha. 1 :350O()0 series (Geol. Sun. S. Au*a.. 
Adelaide). 

r.'iirasTii rn i« k. \_. dvm.\. i ?\. &vONDfcBBQflau ,c 

(.1945) Sequence Mralij.*raphy and (lie interpretation ol 
NeopnUt'OVnic L'iiitll Insliny. I'rrc.tnthtidit AYv 73, J » 25 

. Yn-, niN BCtoGM.C.C, cV Di BosA.KA.dvun 

WotkniL 1 h;, potheses lor I lie on can of Hit- Wiumka 
canvnns iNcnprnlero/oic). Soolh Australia. \nt<K M'ttr, 
frj. IW-Ai J l 'v,;; 
( >l '.IS. K. I 1 ( IM7J)<VOPI 1^ ftrottl Allflrfillifl lsplauafoiy 
N.nes, l:35tl 000 pJMlu^iCul seiics (Ceol Sinv. S. Alls!.. 
Adelaide* 

U'lllAM!/,. K, C., »*KAVvlOKI>. A. R.. C VMI'WA. It 

\ I'iiaii ink. I). (I L '(".9> Momil rainier Piio'litce 
OoOlO^U'ftl Ml:... Sp (1 .;il Series, I £50 000, ((Vol. Sun. 
S Aum,. Adelaide). 

t V Pioiv.. W, V (1987) Stratigraphy of the 



I mktatanu (at. up in Prciv.. W V. (fompllfifl T| k 
Adelaide Gcnsyni-lliie late I'rotern/nie sitali^rapliy. 
M-dinienialuui. palaeontology and ieU"imv" Ret// n , r .■ , 
Y Aiut. Hull. 53. \2S-2iN. 

V\ I'KW. .1. R il%4t TiansgiesMous and rcgmsMPift pp 
175-203 ft Mdiei. K. r_ iio,. "Hrtpcn If) Muriw 

Gfl ,d,cv ' I Macrmllan , New York). 

i >-,i •■ \mn< u t . R. A .l')itr-!N'.>\. .1. I., i IV64) Wiiprnu Gnnip 

gltag ("<''- V.'/rv, (i,,tL \tin: S <\,t\i 9; P- 1 \ 

& . I l%ol PARACHII ISA map sheei. 

Geological Adas of South Ausiiaha. I 250 000 scriwj 

KiCrtl Sitrv S AukI , Adelaide* 
i '■! i- mh i . R \V. i \'>'t2) Tidal dcpositioual .systems pp. 

I^WIH to Walker. K, (J. ,V .lames. V R fHda) I m u 

model* tespitnse to sea level chany/" t( ieoloei' al 
\-Aoeialiou nft aiiada. OrlUirid). 
|>| Bona. |> A. & YWi Di i< lio^ n. C C il'^.i 

Si^l|iiicnl;iry licuIouv and evolution ul an UUtX'tVPpiftN 

shell nuinjiii delta- I .ale I'nilem/iMe Wiwntka [Vainalion. 

Sonlh AnMrati;' Awf/; Aa«W'. ft'l Oi'oi, Unit, 11, %} 

1)79 
IHibN. I A (l'W:al GculO^ n| Hie I Ipalmna Miapir. 
■ nu al hlmdeis Rainies. QtSut't, Lh'ltl \t>h.\. (fail Xltn 

i I'HPh) Sirali^raphie nomenelaiure and 

••eminence siiaUj'iapliy of die kWr WUp&mi Gnmp. 
Adrlatdf ( lensvncliiv: the Sandison Suliuiimp. fin, I 

m 7 im 

i lv;S, a moilet for Motni '.edinieniaiion lioni 
the NL't»pr.aeie./oie Bfftchlnfl formal iuti, Adelaide 
rR-.v.viiilitu- MMJ&UHUlIZi. 51-riS. 

f I VM6a i 1-^LeusM'iial lectouiLs. dtapirisin ,nn\ 
i out tain Mi of seoneiiee boundaries in Neiipioliao/oic 
MiLeev^iouv. ul the Adelaide u'M^.yncline. StHitll 

■Nuuralia. fsth Australian fyohijiuytl Cflnwnttoi), 

■ wk NV> U "i CiValuJit tit v '" n'i\ --,' \tixtmliii 



(|0%h) Stratigraphy ol the KUITfl and 
I mheratana Ciroups in die Willippa Antieline, eenlral 
KhndeiK Ranees, (HuinXwi N<>ics, Grul. Sim, ,Y W/w 
12*) 10 M>. 

I l*) l >(K') A ease loi the falling slaye sysitfiflH iraet 

the significance of forced regreshiv^ de^oMts in die 

NeopltHeHvoit Sandixim Snhyroiip, I <dt A'lMrnfiau 
Geott»Mtctf} Comrtition, Cunht'im. ACT. Li<*>lfyuut 
Sot fury of AiiMKiltu Ab\im i.\ 41, l ^V 

( l^ful) Tin- st-inifieanee ol a lo^ stand 



prngraOiflg wedge hounded by deep water taiKmates ill 
the CaiXlbflftll Milendelhi I iinesiune, KauinanhH. 
Trough. South Ausiralia. Ibut. IJ4, 

& Von Diu IJokOKC C-I,I4K6>A held ya.<le to 
the ucoltmy ei (he late Pieeambrian Wilpina Group. 
Ililleti Cove. Snuih Australia pp, 17-1(1 /// f'arket. A. .1. 
(hil.) Kighlh Australian (jeoloeiVal < onveulion. Due 
day geoIujjit'Hl eveurann-. of iIil- Adi lai.le Kginil ' [( W)\ 
Soc. Aust.. Adelaide), 

& ( ! l ) L M) Sei|uenee sh atiijiapliy ol an 

uui^eil vallev Till the Ncopio(eio/oic Seaelitl 
Sandstone. Adelaide ( ieo-.ynclme. South Au.siraha /// 
Uilryntple. ft-., Zaillliv B & Koyd. R. (I:d^.) "fnii-.ed 
valley systems: origin and '.edimeiitiay se^uenees," So. 

Eiwn, Ptileon. Mim-ml Spn. /',//< 31, MW 222 
lAuan A I\ (15&3J Transi-'ressOL uvu m\v (IK) 
sequence analysis uf the luiasMe MieeesMnn III die 
Sveidiup Kasiif. Canadian Aaiu An hiprla-«> I mi, Jour. 
lAinhS.i.M), V)\ 320. 

"in ■■his. n, (., & Riaiv.. w. v. HV87J SlmOgniphy whhw 

Wilpina CiTfiXipttt Preish, \Y V Uompden 'Tlie AOclaide 
Cietisyneline laie I'ruiero/oie >lru(i^taph\.. sedi 
mentabon. palaeonloK)gv and tectonics." Hull. (<'•■<>! 
Stent $, Mm 5f, Ji 1-254 

i lo miv V, A, L Y Jinkiss, R | f. i ]0X3» Sedimeiilatinii ol 
llie early hdiaearan. Flindi-rs Ran^e^. Soaih Ausiralia, 
Oval Soc, Am. Atm 10. I06-II9T. 

Mom:, I'. \V. il')SS) Stoim dominaled mixed carbonate 
/siliciclaslic shelf sequence displasm^ eyele-. *>t 
laimmoLkv lOiss-siiabtieahnii. laic RrtHcro/oic W'ouoka 
fMMntaliivn. South Australia. SnJ. GwL 58. 2M-254. 

(I00IH A late I'rotero/otc Moim-dominated 
carbnnaie shell .-.eottence: J he Wunnkil KoiTOflUmi m the 
central and sOQlhtTV) llinders Ranyes. Soiah Anvhaha In 
Jilgflf. .1. H <V_ Mnoie. P. S, il.ds) '"The evolution ol' a \nit 
hti a i ill trial i Karlv I'alaco/oie Rill C'limples ||ic 
Adelaide ( Ieo-.\ neline " t'nni Sub \UN> SpiH*> Plifr (<»■ 
177 \<)X 

li:.vi- , R. I. !• | LTOJ I'av 3 Ce.necpf. p| i:dinearan ,u)d 
Ediaciiri;m Ssstems hi Jenkins, R, I, I,, land^ay. I I- .\' 
Walter. M. R, tltdsl 'irckl Guide it; llii' Adelaide 
Cleosvncliite antl AmadetiK flasitv" \uw (h;>i Sun. 
Oft Ciit/hrro. Kept. IVQ.V35. 707, 

lull,/, U K HTtpM, M N. \l Niv>n. I, (i M'"''o 
ANDAMOOKA map Uieei. (,eolo : jieai AtlOK (Jl South 
AnM.alia I ^M) 1)0(1 seric. (Ccol Sur % S, -ai.i, 
Adelaide). 

I 1 1 son, H, & Niv.r-., I i. i looco Mellana map .Ik.i. 
t ...sdorieal Ada-, "I Souih Australia. \lf\% HiO xCtu 
t( i. ol Stin S Aust- Adelaidet. 



AKUHNA AND DEPOT SPRINGS SUBGROUPS 



i 15 



McGinnis, J. P., Driscoll. N. W., Karnlr, G. D.. 
Brumbaugh, W. I), & Camlron, N. ( f 993) Fle\ural 
response of passive margins to deep-sea erosion and 
slope relreal: Implications lor relative sea-level change. 
Geology 21. 643-640. 

Mitciium, R. Mi (l°77) Seismic stratigraphy and global 
changes of sea level. Part I: glossary ot terms used in 
seismic stratigraphy hi Payton, C. E. (Ed.) "Seismic 
Stratigraphy Applications to Hydrocarbon 

Exploration." Aiticr. Assoc, Petrol. GcoL Mem. 26, 205- 
212. 

& Van Wagoner, J. C. (1991) High-frequency 



sequences and their slacking patterns; sequence 
Mrutigraphic evidence of high frequency enstatic cycles. 
Sed. Geol. Id, 131-160. 

. Sancirec, J. B„ Vail, P. R. & Wornardt, W. W. 



(199^) Recognizing sequences and systems tracts from 
well logs, seismic data, and bioslraligraphy: examples 
from the lale Ccno/.ok ollhe Gulf of Mexico lit VVeimer. 
P. & Posamenliei. II. Vv'. (Eds) "Siliciclastie Sequence 
Suaimraphy." Amer. Assoc, Petrol. Geol. Mem. 58, 163- 
197. " 

Pi i mmir, P T S. (1978) Stratigraphy ol the lower Wilpena 
Group (laic Precambrian), Flinders Ranges. South 
Australia. Turns. #, Soc. S. Atist. 102, 25-38, 



Pki-iss, W. V. (1987a) Stratigraphk nomenclafure and 
classification hi Preiss, W. V. (Compiler) "The Adelaide 
Geosyneiinc - late Proterozoic stratigraphy, 
sedimentation, palaeontology and tectonics."* Geol. Sttiv. 
5. Aitst. BulL 53, 29-K4. 

( 1987b) Tectonics of the Adelaide Geosyncline. 

told, 255-282. 

Salvador A. (1994) "International Slraligraphie Guide: A 
guide to slraligraphie classification, terminology and 
procedure" (titter, Union Geol. Sci.. Trondheim & Geol. 
Soc. Amer., Boulder) 2nd edn. 

Van Wagonkr, .1. C„ Muchum. R. M. & Posami N ru;ii, H. 
W. (1987) Seismic stratigraphy using sequence 
stratigraphy, part 2: key definitions ot sequence 
stratigraphy In Bally. A. W, (Ed.) ■■Volume 1. Atlas of 
seismic straligraphy." Amer. Assoc, Petrol. Geol., Studies 
in Geology 27, I 1-14. 

Von Dhr BoRni. C, (\. Ciirisiil-Bi kk, N. & Grady, A, E. 
( 1988) Dcposilionul sequence analysis applied to the late 
Proleiozoie Wilpena Group, Adelaide Geosyncline. Ausi. 
Jour. Earth Sci. $5. 59 7 I , 

Wai.kir, R. G. (1990) Perspective; facies modeling and 
■sequence stratigraphy. Join: Sed. Petrology 60, 777-786. 

Williams. G. E. (1479) Seduneniology. stable-isotope 
geochemistry and palaeoenvironmenl of dolostones 
capping lale Precambrian glacial sequences in Australia, 
Jour, Genl, Soc. Aiest. 26. 377-386. 



STRATIGRAPHY OF THE NEOPROTEROZOIC TENT HILL 

FORMATION AND SIMMENS QUARTZITE AT SOUTH TENT 

HILL ON THE STUART SHELF, SOUTH AUSTRALIA 



By Ian A, Dyson* 



Summary 

Dyson, I. A. (1996) Stratigraphy Of the Neoproterozoic Tent Hill Formation and 
Simmens Quartzite at South Tent Hill on the Stuart Shelf, South Australia. Trans. R. 
Soc. S. Aust. 120(3), 117-129, 29 November, 1996. 

The Tent Hill Formation and Simmens Quartzite represent regressive, shallow marine 
sedimentation of the Sandison Subgroup on the Stuart Shelf. At South Tent Hill, the 
Tent Hill Formation comprises the Tregolana Shale, Lincoln Gap Siltstone and 
Corraberra Sandstone members and represents deposition in a storm-dominated shelf 
environment. Sharp-based, swaley cross-stratified sandstone beds of the Corraberra 
Sandstone Member are interpreted as forced regressive deposits formed above 
fairweather wave base. The overlying Simmens Quartzite was deposited on a broad, 
open shelf that was conducive to tidal amplification. These units are correlated with 
their stratigraphic equivalents in the Adelaide Geosyncline. 

Key Words: Stratigraphy, Neoproterozoic, Tent Hill Formation, Simmens Quartzite, 
Lincoln Gap Siltstone Member, Sandison Subgroup, Stuart Shelf, Adelaide 
Geosyncline. 



TtilHXWttms "I tin' HttVt& \ivnvi i>j V, An\< t iVJftf, 120(^1, I f / P'. 

STRATIGRAPHY OF THE NEOPROTEROZOIC TEN! HILL FORMATION AND 
SIMMENS QIARI Zl IB AT SOUTH TENT HILL ON THE STL/ART SHELF, 

SOUTH AUSTRALIA 

hy Ian A. Dyson 1 ' 

Summary 

1)vso\ I. A. ( l l >%) SuuiigMphv ol the Neoproieri//oic Teni Hill IA)rni;mon and Simmons Quart/he at South 
Tcni Millnn theSiuan Shell", Smith Australia. Trans. &$oc. S. AitsL \2iH^h I 17 \2KK 29 Novcmher, IW&. 

Thi* Tent Hill formation ami Sunmcns Ouarl/ile represent regftUiXtvf?, shnllmv marine seilimeniainiM tij lIlQ 
Samhson Subgiuitp on the Stuart Shelf, Al South Tent Hill, the Tern Hill Formation comprises the TrCgOliimi 
Miiilc. Lincoln Gap Siltsltine tfrtd Cuoabena SunuMiHle members and represents deposition ill a storm 
dominated shch environment Sharp- based, fwiV tfc} emss-^irjtiihrd -sandstone beds of Hie r'orniben:i Sandstone 
Member arv interpreted as forced regressive deposits formed uEw$ fairwcathci wave base The overlying 
Simmens Quart/He was deposited Bit a broad- open shelf thai was conducive ro tidal amplification I heve nuns 
are correlated with their straiipraphic equivalents in the Adelaide Cieosyncliiie. 

KlV WurthM Stratigraphy, Neopiolcro-aMC. Tent Hill Fwrinuliun. S-iiimens Cju;nUi1c, Lincoln Gffip Sdtstoiw- 
Menibei, Sandison Subgroup, Stuail Shelf. Adelaide Geosyneline. 



Introduction 

< hi the Stuart Shelf, the flat lying sediments of the 
Tent Mill Formation crop out west of llie Juntos 
Hinge /one and Adelaide Gcosyneline in South 
Australia. Hie lent Mill Formation t Blown JS85] 
was named after the flnt-topped hills 2? km 
northwest of Port Augusta (Fig- 1 1 As pari ol'a major 
study (Dyson I 1 -)')? 1 ), ihc scdtmenlology and 
stratigraphy- ol the Tent Hill Formation and Sin- mens 
Quart/ite were investigated in the type section at 
Soulh Teni Hill (Fig. 2) and represent the first 
detailed synthesis of ihe sedirnentolngy and 
stratigraphy uf these formations. This paper revises 
the siraiigraphie noinenclaiure for ihc 'lent Hill 
Ruination on ihc Stuarl Shell. 

At Soulh Ten! Mill, the Tent Mil! Formation was 
formally defined as consisting ol three members,. 
namely the Tregolana Shale Member, the Corraberru 
SiOoKtonc Member and the Simmens Qiuur/ii.c 
Member (Dalgarno tri al. I%8». This study has 
elevated the Simmens Quart/ite Member lo 
totmntion status atttl redefined the Curruhciia 
Sandstone Mcmbei, thereby incoiporalmg the 
I iueoln Gap Sillstone Memher into the Tent Hill 
Formation (Fig. %\ The Tent Hill Formation is 
correlative with the Kiachina Formation it the 
Adelaide Gcosyneline. Similarly, ihe overly me 
Simmens Quart/ite may be correlated with the ABC 
Kanye Qunil/iie, Ihe name "I incoln Gap Sihstone 



N.HMMiat renin I'M Klh'kimt t*ealf)$> itfUl Q^ijUtyMCs. 

I iiivoi-)I\ ot Adelaide AdeluiuV S. ,ViM S0U.\ 
|)\Mr (A (PJWyj KedMnetituln«v ;onl .sUiiUs-rriphy *it Hit: 
Ncoprud-Hvtm.' Smidivm Siilv'iunp; u •auirn-dunmtiited shallow 
manm* •icqiifttei: in UjC Adelaide t IcikvikIiik- Soulli \n-:lr:di.i . 
I'M) thrw.. Hinder*; limvL-rsiry orSnuth Atumdia iimpuli | 



Member" has been reserved by the Central RcgjSTci 
of Aiisttahaii Slratigtapluc Names. 



The seen. in at Suuth Tent Hill (Fig. 4). itngmnllv 
describeil by Thomson fl u ()S|, was proposed by 
Dalgarno tt hi \ 1*968) as the type section Un the 'Tenl 
Hill Formation. The underlying formation was 
referred lo as the "Tregolana shales'' by Miles 
(i*)55). Hie torm "Fineolp Gap JPlftgiSWRiiS^ uas 
proposetl by Miles (l l )55) lor the sandy succession 
of Ihe Tent Hill Formation- Crawford ( W64I referred 
lo the lower pan oi' this una as the Corrahetrn 
Sandstone. Thomson 1 1%5) defmetl the Corraberra 
Sandstone Member and the Simmens Quart/ite 
Membet as eonstituenhs of the Tent Hill Formation. 
Coats i l%5) correlated the Tre\uolnna Shale Member 
and Corraherru Sandstone Member with the 
Bruchina Fo^rmaiion, and the Simmens Ouari/ile 
Member with the ABC Range Quartette. Johns 
( I c '68 ) proposed the lerms "Wotimera Shale 
Member'" and "Arcoona Quart/ite Member" on the 
northern Smart Shell where they were considered 
lateral equivalents oi' the Tregolanu Shale and 
Simmens Quart/ite Members, respectively iCoals 

The straligraphy of the lettl Hill Formaltoii was 
reviewed by Coats (1965). Thomson (l%H Johns 
( 1968) and Forbes Si. Preiss ( 19^7)* \\ was ci)nsidered 
u» be ol Mannoan age by rhornsou i 1965). Howevei, 
the sedimentology ot the Tent Hill Formation has 
never been studied m detail and die stratigraphie 
column of the South Tent Hill section by Thomson 
1 1%^) is the only previous attempt 10 identity and 
document the sedimentary structures. Johns ll°T»K» 
considered that the suite of sedimentary Mtuclures m 



118 



I. A. DYSON 




H«, I. Teclomvsedimentary provinces in South Australia, showing localities of sinitigraphic sections in the Adelaide 
Geosyncline and their relation to other localities on Ihe Stuart Shelf and in the 1 orrens Hin^e Zone (after Dyson 1995'). 



TENT HILL FORMATION AND S1MMHNS QttARTZITE 



II') 




Ft£. 2, View of the type section lor the Tent Hill Formation 
(Dalgjrno ,-t ni. l%K)»n the southern face of SotJlh Teill 
Hill. 



* 



WF^r^r 



Fig. 3. Stratigraphy log oi" the Simmons Quart/ite and Tent 
Hill Formation at South Tent Hill falter Dvson I W5'h 



SOUTH TENT HILL 


Stratigraphy 


Sequence 
Stratigraphy 


n 

O 
OH 

o 

m 

00 

y 

o 
ui 

n 
y. 
< 
to 


Simmens 
Quartzite 


metres 

--<*\ 

200 — 

yy 

150 — 

100 — 
50 — 

o — 


I I I I I I 

f m c vc 
mud sand gravel 


Forced 

Regressive 

Systems 

Tract 

(FRST) 














tidal shelf 








.2 
o 

E 

O 

=E 

a> 

I — 


Corraberra 
Sst Member 




shoreface 


Lincoln Gap 
Siltstone 
Member 








Highstand 

Systems 

Tract 

(HST) 




outer shelf 


I 








Tregolana 
Siltstone 
Member 




















parallel lamination ^-S hummocky cross 

^<c\ ripple cross-bedding ^4& swaley cross-strat 
S^y* trough cross-bedding * * * clasts 


stratification 
iftcation 



1211 



I A. DYSON 




I ig, 4 Geological roiwo'f South Tern Hill 



I I.N I llll 1. IOKMM ION ANI ) SIMMLNS QJ AKT/fH 



L?1 



the lent Hill Formation was indicative pf strone 
current action and shallow water sedimentation. 

Stntltxtaphy 

The former Simmens Quarl/ite Membet ot Ihe 
Tent Hill Formation is raised herein to formation 
status to reflect its regional -dguillcunce. Together, 
the lent Hill l-nrmulion and Simmens QuaiT/iie (Pi». 
3) represent regressive sedimcnlaiion of the Sandison 
Subgroup pn HlC Slnai'l Shelf- The Sandison 
Subgroup is an unconformity -bounded deposiumuil 
sequence in ihe sense of Mitchum { 1977) Do ostone 
Of (he intiliM l\ iu^r NiKci.lL't-n;! tiHin.aiitn dues nut 
crop onl in Ihe Tenl Iliils bill is present in driilcore 
front the Stuart Shell- It is rommoniy about 2-h in 
thick and displays a sharp to gradntional base. Ihe 
\n». i'.iKrna Formation was' deposited below storm 
wave bav: on a combined sequence bourd- 
ar\7transj>rcssivc surface that represents an hiatal 
suilace of terrigenous suuvitiou, Ihe lowei lent Hill 
Formal ion comprises the Tregolana Sha e and 
I im oln Qitp Sihstone members that represent the 
lughstaild systems tract of the Naiidisnn Subgroup, 
An interpreted hilling stage or foiced regress i\e 
systems trael i Dyson IWba). comprising Ihe 
ovei lying Oorrabcrra Sandstone Member oi the lent 
Hill Furmaiion and the Simmcns Quarl/ite, is placed 



between the htghsland iiy$tp[pn Had and Ihe 
sequence bonndaty at the top of the Sandison 
Subgroup (Pie 3}. 

The Sandison Subgroup on the Stuart Shelf and in 
Ihe Adelaide Gcosyucliue (Fig, 5) is unconformablv 
overlain by the Wileolo Sandstone and. together wilh 
the Yarloo Shale, is herein assigned to the Aruhna 
Subgroup (Dyson l<)%b). The Yarloo Shale is m turn 
uneonformably overlain by the Wearing Dolomiie 
and. logclher with Ihe Wonoka Formal km. is 
assigned to the Depot Springs Subgroup iDvson 
IW>b). 

Tent Hill formation 

The Tent Hill Formation, about 200 m thick at 
South lent Hill, \$ an upward-sanding unit consisting 
^\' the Tregoluna Shale Member, the 1 ineoln G&p 
Sillslone Member and the Corrabcrra Sandstone 
Member (Fig. 3). It is gtadabonally overlain by the 
Simmons Quart/lie. 



Trcxcltttia Shale M^Hiho 

The Iregoiana Shale Member consists ol 
laminated to thin-bedded, very fine to liue-gramed, 
dark greyish brown sandstone oilerbedded with 
greyish rid *hale (Pig. 6). It is about 60 m thick 




Fiji. 5. Nomenclature and cnrrvluiion t.t ihe Kamhsnn Subgroup t jitter Dyson |9jJ5*): 



\22 



t A IJYSON 



*:t 



w 




L'T^"~ 



. L, 






Fig, 6. Slkile Mid vci;\ thill l>< thm-heddcd line -iirainL-d 
\att.lslnne. IVr't'uhin,! Shfllc Member ol the lent Hill 
I initiation. 



Individual sandstone beds are tint- based with 
occasional grooves and scratches. Internally, the 
sandstones display parallel lamination and current 
ripple cross-lamination analogous |o the Bouma 
sequence for turbidites and are interpreted as having 
been deposited ai or below storm wave base from 
waning, unidirectional currents of storm origin 
Si It Stone beds are characterised by parallel 
lamination. The thickness and frequency of the 
sandstone beds increases up-seelion as the Tregolana 
Shale Member grades into the Lincoln Gap Sihstonc 
Member. The Tregolana Shale Member was 
deposited below storm wave base and is a lateral 
equivalent of the Moolooloo Sihstonc Member or the 
Brachma Formation. 



Lincoln Gap Stlfxtone Mrtnht'r (new HQW) 

The name for this new member of the Tenl Mill 
Formation is derived from '"Lincoln Gap". 24 km 
south of South Tenl Hill. It resurrects, in part, the 
former "Lincoln Gap Flagstones** of Miles (1955) 
that was previously used to include the sandy 
succession above the Tregolana Shale. Mere, the 
Lincoln Gap Sillstone Member is used to describe 
the lower half of the Corraberra Sandstone Meniher 
dial was originally defined by Thomson t [965). It is 
about 40 m thick and consists o\ interbedded greyish 
red shale and thin to medium-bedded, line-grained 
greyish brown sandstone. Flute casts and scratch 
marks are common at the base of sandstone beds 
suggesting current Lransport to the east (Fig. 7). 
Internally, the sandstones commonly display 
horizontal planar lamination. They are, in places, 
capped by interference ripples or asymmetrical 
ripples with sinuous crests. Sandstones displaying 
planar lamination or hummocky cross-stratification 
(FITS) arc capped by near-symmetrical ripples 
Crests ol" the near- symmetrical ripples show a 



FLUTE CASTS 




WAVE RIPPLE CRESTS 



n = 12 



Fig. 7. rVilucoeurrvnl UuUi for the Linenln Gap Sillslmie 
Meniher 



hexagonal pattern or are straight to wavy with 
tuning-fork bifurcations. 

Planar-laminated sandstone beds capped by current 
ripples are interpreted as Bouma BC sequences and 
suggest deposition at or below storm wave base in a 
current-dominated environment. The lluie casts 
suggest the Influence of unidirectional currents that 
were directed off-shore (Fig. 7). Ripple marks on top 
of these sandstone beds indicate the influence of 
unidirectional and oscillatory eurrents. resulting in 
combined-flow ripples (Fig. 8). The presence ol 




TENT HILL FORMATION AND SIMMENS QUARTZITE 

$ 



123 



el 



m^ \'A 




Fig. H, Combined- (low ripples on top of fine-grained sand- 
stone, Lincoln Gap Siltsione Member. The ripple eresLs 
show imperfect bifurcation and note the presence of 
wrinkle marks. 



HCS and planar lamination indicates that storms 
were responsible for the generation of both 
unidirectional and oscillatory currents. Near- 
symmetrical ripples arc interpreted as wave-forrned 
in origin. The orientation of wave ripples suggests a 
north -south palaeoshoreline, These structures 
suggest deposition above storm wave base but below 










fairweather wave base. The Lincoln Gap Silisione 
Member represents deposition in an environment 
where oscillatory currents were dominant over 
unidirectional currents. It is a lateral equivalent of 
the Mooiillah Siltstone Member of die Brachina 
Formation and is sharply overlain by the Corraberra 
Sandstone Member. 

Conaberm Sandstone Member 

Tine Corraberra Sandstone Member is about 25 m 
thick and consists of greyish red, iron-stained, fine to 
medium-grained sandstone (Fig. 9) interbedded with 
greyish brown shale. The sandstone beds are 
commonly micaceous and display heavy mineral 
lamination, swaley cross-stratification (Fig. 10), 
quasi-planar lamination (Fig. I I) and medium-scale 
cross-bedding. They are, in places, capped by 
symmetrical and asymmetrical ripples, interpreted as 
wave and current ripples, respectively. Glauconite. 
inlraformatioual mud elasts, mud drapes, foreset 
bundles, climbing ripple cross-lamination and 
herringbone cross-lamination are also present. 
Several upward-sanding cycles, commonly about 5 
m thick, are present in the Corraberra Sandstone 
Member where swaley cross-stratified sandstone 
beds are commonly erosive into underlying cycles 
that comprise tidal sand sheets. The Corraberra 
Sandstone Member grades upward into the Simmens 
Quart/He (Figs 3, 9). 

The lamination and cross-stratification styles 
within the Corraberra Sandstone Member suggest 
initial deposition at or above fairweather wave base 
in a shorefaee environment where oscillatory- 
dominant storm currents were operative. Cross- 
bedding near the top of the unit suggests the 
increasing influence of tidal currents. A tidally- 
influenced marine environment is also indicated by 
the presence of glauconite, bipolar cross-lamination 
and foreset bundles. The disconformity at the base of 
the Corraberra Sandstone Member marks the onset 







Fig. 9. Chocolate-brown sandstone of the Corraberra 
Sandstone Member in the centre-foregound, gradation- 
ally overlain by quartzareniles of the Simmens Quart/ire. 



Fig. 10. Medium-grained sandstone of the Corraberra 
Sandstone Member displaying SCS. Hammer for scale. 



121 



L A. DYSON 




Fiji. I I. l^Ua.si-plunai lonviAJUion im inedium-jirHini'd sand 
stone Corraberra Sandstone Mem her. Note presence of 
im\ angle cross-bedding show ing palaeoflow to ihe 
right. Leni cap is S. 1 mm in iJiumclei- 



Fig. 12. Quiiil/.uenitc ol tllC Simmcns Quari/ilc Jisplas inv' 

mvanmetl LTosis-hokling of tidal origin and SCS. Note 
smalKeiilc licrriiiL'hoiK cross -heckling ;j! Ivise of ln-il 



pf forced regression in the Sjmdison Subgroup ;ind 
meire-lhiek, swaley eross-stratilrcd sandstone beds 
arc interpreted as forced regressive deposits (Dyson 
IMS' ,193%}, Upward-shallowing cycles * arc 
interpreted as pyrasec|uences in the terminology of 
Van Wagonci (1085), On llic Stuart Shell l!w 
Corraberra Sandstone Membet represents partial 
laiejal equivalents oflhe lower ABC Range Quari/.iU- 
and upper Urachina FoUTUflJOll Ifoin die Adelaide 
( uMsviichne, The iron-rich laeics arc similar In 
equivalents thai ctop out easl of Ihe Torrcns Hinge 
/one /it Kulparo, Ochre Cove. Hal leu Cove and 
Putiapa Cap (Fig. I ). It is ihouglu thai the iron was 
derived from ciosinn of the Mesoprotero7oie 
Pandurra Formation on the Gawler Cralon. 



Simmons Quurt/itc 

The Suuiueus UuarUile is ahoul 100 m thick and 
consists of grey to greyish-white, line to medium- 
piuinvd. thin to very thick-hedded quart/arcnite. 
These beds contain vanouh clusis of volcanic and 
granitic composition, varying in si/e up to 20 mm. 
Compound cross- bedded sets comprising herringbone 
cross-MiatilicatioiK sigmoidal cross-bedding displaying 
loresel bundles, planar-tabular cross-bedding, shale 
elasts, horizontal planar lamination and minor swalcy 
cros.s-stratit'icauon (SCSI are abundant (Figs JZ 13). 
Large-scale, trough cross-bedding is common (Fig. 14). 

Compound cross-bedded sandstone suggests 
deposition uf sand waves in a tidc-doininalcd 
environment where asymmetry of the dominant and 
subordinate currenLs was pronounced, but bipolar 
currents were significant. The luck of HCS and SCS. 
except near the base of the Simmons Quarmte 
suggests shorefaee deposition above lairweathcr 
Wave bast- where- tidal currents were dominant 




Pig, 13. Compound Lioxs-lvtliling in quan/.arcoue . .t Ihe 
Stmmens Quatl/itc 




Fig. 14. Daugh cross-bedding in uV Simmens Quttltfilc 
Lens cap is 52 mm in diameter 



It:-NT Hll.l. I ORMATION AND SIMMERS QtlARTZITIi 



as 



Palacocurrent data show a strung trend towards the 
east and are interpreted as reflecting progradution t»l 
ebb How lidal sand sheets (Rg, 15). Following Coats 
t 1965). lite Simmeivs Quart/He is correlated will i the 
ABC Range Qiiarl/ue in the Adelaide Geosyncline. 



St(_itu'th-t j bouwkiry 

The upper boundary of the Sunmens Qtiart/ite 
does not crop out at South lent Hill However, south 
of Bill's Lookotn near the north- western side or Lake 
foiivns, ,m erusively based sandstone at the top ol 
the Simmcns Quart/He Crops out poorly where it is 
contormably overlain by moderate brown lo greyish 
green shale of the Vat loo Shale. The metre- thick, 
medium-grained off-white sandstone eontains basal 
elasts v>i' shale, lithic sandstone and occasional 
volennics. Sedimentary structures include trough 
uosvbedding, SOS and symmetrical ripples. 

The erosivcly-based sandstone at the top ill" the 
Simmons Q)uarl/ite is interpreted as having been 
deposited in an esiuarine environment- The Uyle of 
cross-bedding in the lower part oldie unit suggests 
deposition on a lluvially-dominated shore i ace A 
.siorm influence is indicated by die presence of SCS 



CROSS-BEDDING 



n = 120 




CURRENT LINEATION 




fig. 15 Palrteoeiirreni tlaiu fur the Siininuiis QuuiV.iji: 



and the symmetrical ripples are imctpreied as having 
been formed by wave action, This unil is U possible 
equivalent of the Wilcolo Sandstone lhat overlies the 
ABC Range Quari/.ite in the Adelaide Gcosyucliile 
where it marks the development of broad (e. 10-20 
km) tttC sed valley (ills lhat in places attain a 
thickness ol some 25-50 m in outcrop They consist 
pf a basal. trough cross-bedded lack's Of fluvial 
origin, overlain by SCS shorel'ace sands. The SCS 
shoreface sands pass rapidly Upward into basmul 
shale of ihe Bunyeroo Formation. The Wilcolo 
Sandstone and Bunyeroo Formation together 
conslilule the Aiuhna Subgroup (Dyson ]99(>b}. Its 
upper boundary is represented by Ihe mavimum 
flooding surface at the base ol the Wearing Dolomite 
which is coincident with the development ol 
kiiomelre-deep canyons previously assigned 10 the 
overlvint! Wonoka Formation (Dyson 1995 , |996bh 



Stratigraphic equivalents of the lent Hill 
1 onnation and Sinmicns Quart/ite 

The redeHned Tent Hill Formation ami Simmens 
Qnarl/ite may be considered partial lateral 
equivalents of the Brachinu Formation. ABC Range 
Quart/he and Ulupa Sihstonc in the Adelaide 
Geosyncline (Fig, 5). The Braehinu Formation was 
dellned by Dalgarno & Johnson ( 1964) as the thick 
succession of sihstoue contormably overlying the 
Nuccalceua Formation and passing upwards into the 
ABC Range Quart/ite tMawson 1939). Together 
with the Seaelilf Sandstone and Nuiealerii.i 
Formation. Ihe Braehinu Formation and ABC Ran^c 
Quart/tie were incorporated into the Sandison 
Subgroup (Dyson 1992). The Bruehma Formation 
and ABC Range Quart/ile crop out at several 
localities within the Mount Lolly and Flinders 
ranges, of which the latter occurrences appear lo 
display lateral continuity. A study by Dyson l|992 v 
I995 1 ) focused on well-exposed sections at Hallett 
Cove. Ochre Cove. Mount Terrible, Kulpara, Hidden 
Gorge. Wyaeea Bluff. Partacooua wtKi Bunyeroo 
Gorge (Fig. I ), The Ulupa Siltslone is best developed 
lo the easl and northeast parts of Ihe Adelaide 
Geosyncline on the BCRRA. COPLFY. CLARY, 
ORROROO and MARRFF. 1:250 000 geological 
sheets. The Braehina Formation and overlying ABC 
Range C)uart/ite constitute an overall upward 
sanding succession. They lepiesenl regressive 
sedimentation of the Sandison Subgroup and arc 
therefore defined as all the strata overlying the 
maximum Hooding surface/down lap surface, 
icpresenled by the Nuccaleemi Formation, lo die 
sequence boundary at the base of the Aruhna 
Subgroup. 



I Hi 



I \ l)VSM\ 



ftnnhiHtf ffartmtTfim 

ITio Una liiim InniKitioi) comprises lout inembers_ 
Kidi td which can he dclincd by it- HlbofuCiWs < I-iv 

5j lw....f il.i. i i ficfj the Bttylev Range Siiistont 
i [flK'i and the C'oiraberra Sandstone Member, aiv 
.'"ti%i\l'.ied lateral equivalents (Dyson |q*)Vi The 
Bnylcy Range Silicone Member docs tlijl crop o Lit at 
llallctl Onv 01 Ktilparn (Fig. I), The ( 'oiTubcnii 
Sandstone Member ol tin.' le.a Mill formation thai 
«.ni|.-. uui uii (he Stuart Shell has nn equivaleni thai is 

included in Ihc Rnwhinn Formation m Kallen Cow 

iI\vmiii I'm 1 ), The hthofaeics Ml 1 ihc Braehinn 
I 01 malum a I Hallctl Cove display .< suite ul 
M.limcuiarv •anuniir-. frUggtSStftei ol deposition 
under the influence of greal storms- These siruunre;-. 
including accompanying pulaeucuroeiit data, aie 
illustrated and described more Tullv by Pyson 
<IW, IW5>_ 

AIU' Range OuitiizUe 

The ABC Range Quart/He is eomposeil of pale 
pmkish gfeji lo greyish white, thief bedded to 
massive, fine to medium-grained, slightly fLfdspaliue 

i] "'• Mid tumor iiHerhedded shale and One 

gi aiiieU sandstone. It is characterised hy ihe 
ahundance of planar-tabular, hemnghonc and large 
seale trough cross-bedding, asymmetrical ripples 
wiih sinuous crests. fla«J and leiUieular bedding, 
mudciacks, mud drapes ami mud inlraclasls. 
Slraigbl-crcsted symmetrical ripples also cap some 
sandstone beds. Compound ems -.-bedded sets consist 
ol small-scale cross bedding separated hy inclined, 
• •-i.rnvr set boundaries. Afl Ihe inclination ol Ihe 
itiaMci heddiny planes decreases, Ihe hipolariry o| 
III.- cross bedding becomes more prevalent 
Mcganpplc cross- hedded sets, commonly URJ- 
dircciional. display a ihinthid-. alternation ol sandy 
n-iesct bundles, hounded by reactivation surlaees, mud 
ouplets and/or mud drupes. Sandstone channels are 
characterised hy gently inclined lateral accretion 
surfaces and low angle (ruination surlaees. SCS and 
Intii/ontiil-phinar lamination are occasionally 
observed near the base or some minor qujrt/uc 
anils, MCS is resirieted (n intervals ol inlerbedded 
shale and linn to medium bedded fine sandstone ami 
quatt/ilc. Thus, Ihe suite ol sedimentary sirueiures hi 
Ihe .ABC Range Quart/ile described Korc suggests 
dial ii was deposited in a shallow marine environment 
where lalal currents were domuiani over slorm and 
WOVC action, 

/ )ft/fVI SillKtiOlt 

The Clupa Siltstone e- a regionally significant unit 
that can be mapped over a large area of the Adelaide 
Ceosynclinc The type section of the UJllpa Sihslone 
was defined by Mirams t !%-|) near Moum Bryan on 
HURRA where it was described as a succession ol 



einceu grey ^nd locally pmple shales. Phunmer 
1 197^ Recognised the three members ol the BttloKiltiJ 
Ponnnliuc in ihe I'lupa Siltstone at Oodlii Wina 
<figs I. 5) and recommended the icrm "I llipll 
SilKM.nc he relinquished. However, roibcs A. Piens 
L-1U87] proposed ivu-uiiiHi |>i 1 1 H.- -.tratu'raphic niiP.c 
Ii.t.ukv ihtj aigued lhal tee.ional mappabiliiy ryfihe 
con itrliicnJ incmhers had not been estalibshed 
tiltK.la. a s of the Ulupil Sihslone are disi:il 
equiviilents ol the coiiMiluenl members of \\l< 
I : irai'ton.» Pi)| m^'hm and ure iherclore Iiiik 
"h. .j:*-,mVc. 'Ihe Ukipa Siltstone rcpieseiii> 
" ii ai\c -.hallow marine sedimentaiioiL on the 
middle to outer shell. Palaeocurreut data show i 
v,idc range ol eurreul directica^ (flys*iu pi'is | 
sueg.-stmg an ..•.Kiri)iimeMi where unidirectional and 
o%eillat»*ry currents were interactive. 

Depositionul mudel rur the lent Hill lnriuatinn 
and Siuiniens Quartzite 

The Simmcns Quurt/ite and constituent membcis 
ol' the Tent Mill fiumalion may be defined by llicir 
liihotaeics. Ihe Tent Mill I ormation is an upwurd- 
sanding sueccssu)n ol irilethedded shale and flhe 
LMuiued sandstone. Sandstone beds display several 
sedimentary structures associated w ith -lotn, 
deposition such as BC liouma sequences. HCS. 
micio-HC^S and quasi-planar luminalion. Ihe 
'Trecohuia Shale Member was deposited below Mucin 
wave base, Willi progressive shallow ing above stonn 
wave base, intcrbedded shale and saudMonc ol the 
l.incivln dap Siltstone Member were deposited M 
the top of this succession, die Corritheroi Sandston c 
Member consists of several sharp-based shorela.. 
sandstones about 1-2 m thick thai ilispl.tv SC "s 
These SCS .sandstones :iiv lelerred CO U> atlachvd 
shorelaee deposits. Some SCS >niKKP<nc-. ai- 
complclely enclosed within shale and are relerrc.l 111 
as delached shoief.n.c depcvsit;-, The base of ejdi 
S(JS sandstone is a liigli-frvqucncy M*qaenee 
boundats 1 The erosive shear face deiwrats „.k 
iiiierpreled as kneed n.virc^iVL tJcjlO^ftfi The 
Conaberra Sandstone Mcaubcr was deposile<l al>ovc 
lah weather wave base in an environment lhat WtM 
storm Jominaled bill where tidal »ieti\ity was aJsu 
sigmncaut, U is grjdaiionally overkiin by p n 
sequences ot tide-dominated tiuaii/aiemies and 
llthtfi sandstones ol the Simmcns Qunrl/ile. The 
Siiiunens Qtiart/itc represents continued shallowing 
of the sea in which the Tent Hill and Hraelima 
formations wen- deposucd. Combined with a high 
sediment supply, proyradation of Ihe lower shoreface 
resulted in a wide, shalhiw shell' which was 
conducive to tidal amplification. No submarine fan 
deposits have been recognised at this siratigraphi. 
level elsewhere m l.he basin 



II, Nl Itll I, formation AM».siMMi:N\snu,\in/ni 



.:- 



The Ten I Hill Formation inul Simmcns Ouaiuilc 
represent pro-gradation ol a lide-dormnalcd shorelace 
ill-Ili a storm-dominated, shallow shelf cm ironmenL 
T|ns regressive succession is composed of a number 
.'I upward sanding cycles '|i t n M CpAlClfrQd fc'ithJOtf 
hierarchy ol iransgressiyr-te^resvivc cycles, TIhsc 
ivvics .tic initially aggradational in cfuuacLei and 
beutirue increasingly progiadatiunat upscuion. A 
very thick of flap wedge developed rrurfl litis 
pro>jt.Hl,i!H»ii mid gravitational instability resulted in 
e\lenstonal lauding and enhanced ;UbS[(lC4li r 
Palaeocurrenl data suggest a noillvsoulh, 'idally 
inllneneed shoreline. vSediineni was derived horn the 
tiavelei ( raioii lo I he west and transported eastwards. 
The deposjlional environment shallowed to flbOVC 
lau weather wave base across a relatively narrow 
shell and sediment progiadcd acipsa the Toncns 
Hinge /one into ihe Adelaide Geosynelinc. Here. 
thickness of die Sandismi Subgroup w, t s affected by 
syn-deposilional lcclomes. Coals (l%5) suyeeMcLl 
thai Ihe overlying Won ok a formation and Pound 
Snheioiip Here nni deposited on the Smart Shell. Inn. 
were re aricled lo (he Adelaide Cleosyncline because 
'tlsyn deposilnMial faidling across the Timci'i^ Hinge 
'/one. The urcuale Irend icpicsented by the Tmren> 
llm^e /one marked (he possible L-dgc nf ihe tormer 
filiclf during deposition of the lower Woimka 
I oimalion. However. ulentirieaLion or ihe Wonoka 
I ormalinii m dnllcmc PfOTll ihe eastern Stiuil Shelf 
iV.e . fjopccchce 2» suggests that a major trans- 
gic%su<n iicemied across ihe Toircns Hinge Zone at 
ibis lime. Deposition ol ihick P.unyeroo sediments in 
ihe Adelaide Geosynelinc v\u>. contemporaneous 
with active subsidence. Regional instability 
contributed lo the incision ol" VVonoka ianyons on die 
western edge Cll ihe (ieosy ucline. 

Discussion 

I oh id n vrc\n * depi-'sns 

In the classic hwon sequence sonographic model, 
Ihe Type J deposition;)! sequence cons sis ol 
lnvss|;md. trunsgrvsstvc -^ Inch-.iaiui -.vsterns duels 
which an- sihciiutticalK lied lo specific increments 
ol'ihc eustalic curve. Hmvesei, an increasing \o|nnv 
of lileraliirc suggests Ihal deposition during a dative 
fall in sea level moy be placed into a rourth systems 
trad between the highland systems tract fBST) tuul 
the .cgiienee boundary. This systems Iruel has been 
previously referred lo as the falling slage or forced 
regressive systems iiacl (e.g.. HumI ..V tucker |9$2) T 
Study ol progradahonal tidal sand sheets and shaiy- 
bascd shoreface deposits of IheTenl Hill Foimalion 
and Simmcns Quart/ite at South Tent Hill, and 
similar deposits of the Brachina Formation and AEfC 
Range Quart/ilc at llallctt Cove. Kulpara. Bunyeroo 
Gorge anil Trebilcock CJap ( Fig. I ) suggests Ihal lliey 



may be ;<ssigned Ul (he lulling stage system^ ttuci 
(Hyson i nu n;it. I he-e Minis represent irgnM^t 
sedimenlalion ol Iht Sandison Subgroup •>!» 'he 
Si] 'in Shelf and in the Adelaide (tei.sym line. The 
lower a\\i\ upper boundaries ol the the Tailing stage 
systems Had (FSST) UW flXpd e.n Ok relauve sea 
level curve, However, the ineremenls n| Mic OXllOf 
sysienis iracls are not fixed and will vary due 10 
subsidence raie and sediiueui Supply. Shoietaee 
sandstone displaying SCS ai ihe base d1 the 
i i».r ( hi!na Samlsloiu- Membei convspunds to tbe 
ba i -.' ol Ihe l : SST. Its oppci -boundary is ibr ■ l r..,uciicc 
boundary which is delinai here- as ihe lowest point (if 
relauve sea level. Ihe conclaiivc eoni.-nnity may he 
auakk yon, to |he tkiwnlap -urlacc .a discora^miiiy a\ 
Ihe top k>\ submarine l.-ns m earlier p^mTii m'>dcU |i 
passes epdip into the subacnal uiiconbainiiy 
associated with ihe sequence lioundary 



Sfh'ij f <i\thj\nii v ittitl i'iiliiit'4 •unvim 

ShMo-ta^e siiuui drp.^Hs fif fhe < 'm uilvi -',. 
Sandstone Mcrnbei vveic p<t.Nibly ■.lepivsited j|1 a 
mcsolidal enviroiinieni Willi a ndal tangeol some 2- 
4 m. Such vn\ itiauiicno. ate siorui or nave 
dominjlLd i l.^alry tuple l l » t )2i The Cm rubci r;i 
S-UmI-Umic McmiIui 1$ dhi.Olv sHL.rrOrd by od.il 

sand shcel deposits ol Ihe ARC Range Qunrt/iic tn 
the southern part of ihe '\dckuile (iecr^viicliue. This 
Miggesis ihal lidal overprmting oi storm and wave 
iTlrcis extended well nut acrov. ttte upper •diorefacc 
The relalive inllucnce of stoiins tlccieased a* llie 
Lidal curreni speeds iiKa*ased so ihaL clislal pail.^ o\ 
the sand -heel contained storm-genetatcd Htntcmres. 
The siorm-dounuaicd -hoivlace system was replaced 
by prograding lido dominated delms and 0[>eu coast 
tidal flats. Mud \va^ deposited bcyi'lid ihe depth anil 
mnije of tidal reworking, 'lidal ehtinnels wiihm the 
Simmcns Quart/he weie possibly hieised to shallow 
sublidal depths on the shorelace. based t»n the rarity 
ul S(V The depth of incision suggests a I > t g» h 
mesutida! lo possible macrobdal tango along the 
palacushorclities During deposition of ihe KSS I. lidal 
rantie may have been limited due lo ihe relative kill 
in sea level- 
Wave ripple orientations su^ecsi a regional north 
south shoreline C la-tic material was soureed from 
the west. Asymmetry of the tidal iveime is supported 
by the dominant uiiiniodal trend hn rij»ple cross 
bedding, The wide spicad ol these data suggests that 
longshore currents vvetv operative, Shoaline 
lairwealher and stoitn waves initiated longshore 
currents that transported sand ^ the shoreface. 
rouimly parallel lo the shoreline. Palaeoeurrenl data 
from slnrm-influeneed lithokicies in the Brachiua 
f r ormali'>n at Hallelt C o\e are desctibcd bv l)ysot< 
(l'» t »5 t . IVfcS), 



i:s 



i v in, son 



/ ithf>/iH-ic\ rrlatio/ishipy and cot rcliiiions 

The Corraherra Sandstone Member pf ihe Ton I Hill 
Puirnalion ,n SomhTcnt Hill and on ihc Slliarl Shvlf 
comprises ihc lower part of the sandy succession 
overlying the Tregolana Shale Membei (Crawford 
1964)j Pitholacics resembling tho.se of the 
Conaberra Sandslone Member also crop out at 
Ochre Cove i Dyson I995'l Halleli Cove and 
Kulpara tDyson l l .)92) and al Puttapa Gap near 
Bellaiia (Coats 1 96$), Granular, medium lo coarse- 
grained sandstone and dark red lo reddish brown 
shale al Ochre Coxc (Fig. 1 1, previously identdied as 
Bunycruo Portnalicrn (Dyson P>92). was 
reinterpreted as lithofacies of the Corraberra 
Sandstone Member of Ihc Brachina Formation 
(Dyson P)9V)- At Kulpara. faeics of ihc Corraherra 
Sandstone Member are inteibeddcd with the lower 
\PC Range Quart/ite The Roy ley Ranyc -Silicone 
Member CtOpi uUl north o\' Pichi Richi Pass at 
Middle Gorge. Partacoona, Warrakimbo Gorge. 
Rluck Gap. Bunycroo Gorge. Brachina Gorge and 
I'inke Springs (Fig. |), SouLh of Partacoona and 
adjacent to the Torrens Hinge Zone, the storm- 
dominated faeics ol die Conaberra Sandslonc 
Membei are predominant. This suggests that the 
< "(tnaherra Sandstone and Bayley Range Sillstonc 
members, both deposited above fairwealher wave 
base, are lateral equivalents, Ptirtheimore, they arc 
partial lateral equivalents ol the lower ABC Range 
Quan/.ile, 



The terms l, SIntmoiia Quart/ite" and "ABC Ranee 
Quart/ite" are used herein lo describe silica- 
cemented quart/ite or orth<H|ituil/iie in which the 
dominant mineralogy is ovci 90'/r quart/,. Pcllijohn 
t>( nL (197?) peeler the use of the term 
"qiwrunrenitc' 4 over 'orihoquart/ile" for those 
sedinieiils in which Ihc derntal traction is $5% or 
more quartz. The ABC 1 Range- Quart/ite has been 
mapped at the first appearance Of thick, laterally 
extensive white quart/ite (e.g.. Webb & von der 
Porch l u f>2: Dalgarno & Johnson 1966). Apparent 
mlertongiiing of the Tent Hill Porrnatmii ami 
Simmons Quart/ite »»n the Stuart Shell, and between 
Ihc Brachina Formation and ABC Range Quart/ite in 
die Adelaide Geosynclinc can be generated by the 
stacking of these lithofacics on a parasequenee scale 
Similarly, interionguing between the constituent 
members ol Ihc Brachina Formation and Tent Hill 
Formation may he explained in this manner 



Acknowledgments 

The study of the lent Hill Formation and Sunmens 
Quari/ite comprised part of a PhD dissertation by ihc 
author at Flinders University. Wolfgang Prciss and an 
anonymous reviewer read the manuscript critically 
ami contributed many helpful suggestions Gail 
Jackson cttld Gha/i Kraishnan dratted the figures 



References 



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country naNscd over from Port Augitsia to Pucla. fori. 

I'np S- AuM. 45 _ 
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Pake fiirrfiis A\eu>. (Jnurr. (hel. Ntty.-i (rrnl Sm\> S 

&tttr. l6,o.( i 
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Si _ _ I l%o) PAKACt III .NA map ^lieet. 

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^et|uenec Mrallgiaphy <A itu FfiWdi VVd[n_-na Gn».i|i, 
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_ (l°9Sj A model for storm •.cdaneni.ilion Prom 

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Neoprotero/oie Surutiswn SilbgrctlJp. Gf*>h)^ual Smn-i\ 

Oj Ati\ttoNti Ahshtu t\ XX I2\ / <th \tttitJVHun 

Gt.>tf\i;ii til ( ami fttintt, Cituhi'fni, ACT 

(J*W» Stiatiytaphv ui ihe Aruluni and Depot 

Springs Suh.eroups. Truttt. R. Sor. S. Aast 120, 101 I l\ 
Fokbis, H G. A; Puuss. W 1 V. tPJN?) SLratigraph> ol ihc 

WdpeiiaGnutp /'/ Pretss. W. V. (entnpilei ) "The Adelaide 

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Surv.XAuM 51,2:1 1-^4 
I li NT I), (t IVCKKU, M. P t, 1992) Slranded parase<)ncnee^ 

and ihe foived regre^ive vsetlye systems Iracl deposiliv'M 

<li.nn- base level gfl|, Sal Qevf. 81. l- l) , 



TENT HILL FORMATION AND SIMMHNS QUARTZITE 



129 



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2 12 



Pettijohn, F. J., Potter, P. E. & Siever, R. (1972) "Sand 

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series (Geol. Surv. S. Aust., Adelaide). 



TRANSACTIONS OF THE 

ROYAL SOCIETY 
OF SOUTH AUSTRALIA 



INCORPORATED 



VOL. 120, PART 4 



BIOLOGY OF THE EUCALYPT GALL-FORMING FLY, 
FERGUSONINA FLAVICORNIS MALLOCH (DIPTERA: 

FERGUSONINIDAE) AND ITS ASSOCIATED 

HYMENOPTERANS IN SOUTH AUSTRALIA, WITH A 

DESCRIPTION OF A NEW SPECIES OF BRACON 

(HYMENOPTERA: BRACONIDAE) 



ByG. 5. Taylor, A D. Austin & K. A. Dawes 2 



Summary 

Taylor, G. S., Austin, A. D. & Davies, K. A. (1996) Biology of the eucalypt gall- 
forming fly Fergusonina flavicornis Malloch (Diptera: Fergusoninidae) and its 
associated hymenopterans in South Australia, with a description of a new species of 
Bracon (Hymenoptera: Braconidae). Trans. R. Soc. S. Aust. 120(4), 131-146, 29 
November, 1996. 

Galls initiated by the association of Fergusonina flavicornis Malloch (Insecta: 
Diptera: Fergusoninidae) and Fergusobia sp. (Nematoda: Sphaerulariidae) on 
Eucalyptus camaldulensis Dehnh. at Goolwa, South Australia were monitored during 
a single, late summer generation for emergence of flies and associated Hymenoptera. 
The morphology of the galls is described for the first time, A total of 12 
hymenopteran species was reared from galls, twice the number previously recorded 
from Fergusonina/Fergusobia galls. There was a strong positive correlation between 
emergence of flies, combined totals of flies and wasps and gall size. Bracon 
fergusoninus sp. nov., a probable primary parasitoid of F. flavicornis larvae, is 
described. Notes on the biology and taxonomy of each hymenopteran species are 
presented along with an illustrated key to their identification. 

Key Words: Eucalyptus, gall-forming, Fergusonina, Diptera, Fergusoninidae, 
parasitic Hymenoptera, Braconidae, Bracon, Fergusobia, Nematoda, Sphaerulariidae. 



TnmxurtfHisof ihr tfmw/.V«Wv flfjfc /Wv/, HWrik 120/ k I3J I >f> 

BIOLOGY OF THE EUCALYPT GALL-FORMING FLY, FEKGUS(h\t,\A 

I [AVICORMS MALLOCH (DIPTERA; FERGUSONINTDAE) AND ITS ASSOCIATED 

IIYMENOPTERANS IN SOUTH AUSTRALIA, WITH A DESCRIPTION OF A NEW 

SPECIES OF ttRACON (HYMENOP I ERA: RKACONIUAE) 

by d. S. Taylor. A. D. Acstin & K. A. Dwn s 1 

Summary 

T\s i ou. & S.. AUMlv. A. D & D wirv K A. i lOj$J Biology <rf the eucalypi gull-forming lly hi^nM>nnm 
flmtiontis Mulloch (D'lpMii: l-eigiisoninidact and iis associated hvinenoptemns in Soulh Australia, with u 
description ol ;i new species ill Hiiian) iHvinenopleta I.Uaeotiidnel irons, R, S'n . A \Uiti 120 (4 k 13 I-I4h. 2*-) 
November. 19%. 

flails initialed by Mil* association ol Ftr^usttniitu ff<i\nt,>tnii Mallodi (Insecla; D'/pterj; 1 -ergiivomuidac) ami 
I i'r#us<>hiit sp. (NoinukxIiV Sphaerulatiidac) on i uralyflHOt cmnndiuhitsis Delmh. at Goolwa. Smith Australia 
ucie mummied duong a single, laie summer generation lor emeniiuec ol fibs and associtiicd Hymenopieia. 
The tnocpbulogg of the galls ifl d^ijbed for l)\c Htai umft. A fmal feJ 1 2 hymenopu-ran species was aaivd ftom 
pills, twice the number previously recorded froni f'('r^it^fni>i'i/hi^)i.\i*hi/t ualK There was a stiong posiiue 
i ■'•nvliiiiun hcTWCcti emergence 1 ol flies, combined totals ol I Hon and wasps and gall !*r*'cr HwpttH <> nmsotmtHs 
<p iiuv.. a probable primary purusUoid ol £ fh<\nt*<tn:< larvae, is described, Noieson the biology and taxonomy 
ol each ItymcuopU'ian species arc prcscniod along with an illustrated key in ibcii idemilieuiion 

Km Whkum EuWil'wtM) gatl-lormmg, Ft'rt>ust/nino> Dipu-i. : . I comsoiimidai', paiasitic llymcmiplern. 
Uraconidai-, linn \m. /w'/^Wtfpfti Ncniatoda, Npliaetuiariidae, 



Introduction 

The biology of gall-associated Hyittenopieru is 
fundamentally complex and it is oil en cxb*einely 
difficult to determine the it tie relationships ol species 
iBoucek I9KS: Gough & MeMahon 1988; Nanmanii 
Wl; Schomoggc ct uL WO), For instance, galls 
may contain a range ol biologically different 
Hyrnenopteta. vizu, primary parasitoids. facultative 
and obligate byperparasiloids and nuiuiliiies. The 
biology v>l (he latter group is particularly difficult to 
unravel. htquihnes arc species lhat live inside galls of 
other ceddogenic insects and are phytophagous lor 
all or some of their larval development. 1'hey often 
kill the primary gall-forming -species at an early stage 
of gall development and then increase the si/i nf the 
gall, thereby making it difficult to determine whether 
nr not tin.* species concerned is indeed Qfl inquilinc ur 
the primary gall former. Other ini|uiliues apparently 
invade galls and k-cd on their tissues without 
disrupting the primary eecidogeuie species. 

One of the M081 complex gall associations known 
is thai between Fcr^nsonitut M'p. tlnsccta; Dtptera: 
Kergiisouinidae) and Fer«tisohia spp. iNeinaloda: 
lylenchida: Sphaeruiariidae) in galls on Myriacc.ic 
(Hshcrei Niekle I96X; Harris \^l: Davies & Lloyd 
I'WVh However, the only detailed study of ilie 



I DepitrlmenL iH Crop Prolet lion, Wane Campus Ihe 
t 'niversilv ol Adelaide PMB I fiteii Osmond S All-sl 



biologv arid phenology oi n^r^usonintt Mutloeh flics 
is that of pifliic (JW7i. wlio conunented on tht* 
possible role* t»! hymeuopterans and other gall 
iiK|Ui lines in tile seasonal fluctuations of '!>' 
populalioiis. Jhls study showed lhat numerous w asps 
were associated with l-crxiiwtimui/f'rrsttstihiti Flower 
hud galls. Although Currie ^ 1 937 1 exammed ihe 
hiolitgy of this systen% the ta.vonomic knowledge Ul 
the lime was limited and the biology of the species 
involved was nol well understood. 

This paper reports on a study of the leaf galls of 
fttrgtWnitttt fltlfhwWh (vtWllOCft associated with an 
undescribed species ol l-t'i^a.snhtti Cuirif ami ihe 
associated guild ol' hymenopterans from galls on 
Fjttnlyputs i'umulihilfn.sis Dehnh. at Cioolwa. South 
Australia. Inlovniatiou is presented on gall 
morphology, the pheiiolonv of the fly and wasps, the 
probable biology of the nymenopteran species 
involved, their taxonomy, and the relationship 
between numbers o\ fhes and wasps and gall size. A 
new species of Bnmm h, thought to be a primary 
parasitoid of Ft'/^usoni/ui. is described and an 
illustrated key to the 12 gall associated hymenop- 
leums is presented, This iohirmation is compared 
wilh Cuiric\-> (1^371 Mud> and is discussed in 
relation to the inherent difllcullies in determining Ihe 
biology of gall associated wasps. 

Materials and Methods 

Seven mature galls on an ornamental, 4 in high /:. 

cam(tf(hil' j "sis sapling at Cioolwa, (35 31 ' S, 



1 .12 



<; S 1/vMwK A |i M'STINtS; K.A. DAVIf.S 



\3K 46' l;> South Auslralia were titged with muslin 
in eatty March. 1995, preceding emergence of adult 
£ fittvivomh and other gall occupants, .in. I 
periodically rUOftllOKCI Until after senescence ol" Ihc 
gull and when Hies unci wasps ceased to emerge, Any 
insects thai had emerged aL each sampling time were 
collected, idcntilied and counted. Gulls were dried 
and weighed to compare gall mass with loial insect 
emergences, For observations kw\ gull development 
and morphology, young fresh galls were sliced open 
in water by u series of truitsvetse euls Abbreviations 
used are: AN It 1 (Australian National Insect 
Collection. CS1RO. Canberra) and WARI (Duncan 
Swan Insect Collection. Waite Campus, The 
University ol Adelaide). Voucher material ol F. 
jluvk-onus has been lodged at ANIC aitd WARI and 
malcrial of all hvmenopferun specie--. is deposited in 
WARI 

Results and Observations 

fytstcihuuoti and host plow h\so<htih*n 

f'trxti.Hotitta flavic<>rni\ was the only di pier an 
reared from an ornamental specimen of /.. 
mntuhttilctt\L\ ai Cioolwa. Similar gulls have heen 
collected by us from several other South Australian 
localities Irom naturally-ocvurmig /. famuhittU'ittts 
neat Verdun and near Mil.ing. and ("torn litis sjirte 
host in Ihe Adehiide suburbs of Hyde Park. Rose 
Park. Tusmore and Crrbrac. Fi*r$ltfOtiifKl jhtviforni\ 
was deserihed from a single female specimen from 
Sydney. NSW (Malioch 1925 1, Larvae, males and 
galls have remained undescribed to date. This is the 
first host plant record ror the species and first 
description o\' the gall {see below). We have also 
collected all lite history stages ol /. fftt\'F<lurtf(i and 
Ihe associated, uudeseribed species o[ i<'iyjt\obiu 
and plan t<> discuss ihcse in ;i separate publication. 

fittll ni(iif)httiit\;x anil jonmaint} 

The galls of /; f(avi<<trnis are formed from 
terminal leaf buds, matme galls (Fig, I) being 
bulbous structures, mostly ovoid m longitudinal 
cross section. IS 50 mm in length by 1 2-20 mm in 
diameter. Mean dry weight of mature galls was 2,22 
± \A5 g (range 84-5. 16 g, n = 7). Calls appear to 
consist of a single leaf bud comprising a number of 
leaves, with Ihe ventral surface of ihe ouleiniost 
foiming the eMeinnl face Of'the gall. The growth of 
these outer leaves sometimes continues unaffected 
beyond Ihe gall proper, cither running along Ihe side 
of Ihe gall or emerging as normal leaf (issue beyond 
its apes. The stem supporting the gull usually 
develops a bend just before its insertion m the bnsc 
of the gall. Transverse sections showed that the 
galled tissue is soft, consisting ol pareneliyrnuton , 
tissue, ahd that h llu\n<>nti\ larvae develop in 



chserclc cavilies. These are oval m longitudinal 
outline and distributed throughout the gall. The 
cavilies ure lined with plant cells winch aie paler in 
colour Hun ihe parenchyma cells between diem. 
I n^ustfhiu nematodes are found in the cavities with 
ihe fly larvais 

Kir the generation of ft jhtvitornis winch began in 
late March, small, recently initiated galls wcic 
observed, tour of these were collected and dissected 
and Ihe development of ethers was monitored during 
the study. Pull gall sr/c was reached on the Live in 
two to four weeU from g;i!l initiation, The followimj 
obscrvalions were made on the dissected galls. The 
smallest gall. H mm in length contained JO spindle- 
shaped h. fhtviiornts eggs (Fig. 2) within which 
developing lai vac could be seen. It contained many 
|i,iri!ienogei>etu fljjj juvenile nematodes and some 
nematode eggs. Al this stage of development no 
c.iv'uie-. were present. Another. Ml mm long. 
contained sever;il eggs, a few first mstnr larvae ol 
fojfW&whtA arul ni, tin parthenogc-netiv n^Dfnilode^ 
and eggs. Some llv larvae were found within 
cavilies, ihe small cavtlies being ringed with tnanv 
white tells. A 12 mm long gall contained firsi ami 
-.ec'ind insiur larvae of h't'i^/isi>nintt, all wilhin 
cavities, ,tm\ many purthenogcnelic nematodes .mm! 
egg- Ihc largest gulf about 30 mm in lenglh. 
contained first and second insiar lly larvae within 




f m i, rerrnurnl letif imd gull nt h i iH*twttttni 

liih'K •itu,vlrt:>.ti\t>hiu on f.mni^mn mii(ihliitrtt\i\ .il 
Citmlwa Souili Australia Soak bill = ^5 nun. 



BfOLfXIY OP FtiNOUStiNtNA AND UN ANSOCIAlf D IIVMI'.MM'TrUA 



131 






- 



J 



Fiy,2. F.gg <A ii'iilHsnntiut }lti\i< omis Irom u tcrmitiiil leaf 

buii gall From l.tunhpms vtimnhhifcnsh at CiutHwa, 
South Ausinilui Settle bar- U.25 mm, 



well-defined cavilie.x. none ol which was ringed wiili 
while culls. |l contained many parlhehogenclic 
female, juvenile and male nematodes mid egg*, The 
ihicc larger gulls cadi contained brown 'atie&kin 
running from Ihc centre 10 il»e amor edges. 
apparently lormed by die darkening 0l fotflCt pfoft< 
cell walls, these observations on the process of girll 
induction indicate thai /• jhivinnins nod nematodes 
arc always present together and dial both fly larvae 
and nematodes continue, lo develop, ihus increasing 
die si/c ol Hie cavities. We have nol been able 10 
determine whether gall initiation requires both 
species to be present or if only one is rci|uired. Given 
dial nematodes are present in the early gall maln\ 
pljoi to hatching of fly Cggii, it is possible tbat 
initially, gall lissuc is produced in ivsponsc to 
nemati'tle m\ iviionx (sec Discussion). 

A; S 'C cinJ (h'fuil \hh'IJ of Ferguxouina flavicornis 

The egg of f. /hiviituvis is spindle-shaped and 
tapers In a shod, blunt 'pedicel' at the micropylar 
tautcrinr) end and to a somewhat lunger, wpwcO 
process at the other end (Fig. 2). The egg is IXfij + 
SI 0.06 mm long hy 0.2o ± SF 0.03 mm wide (n = 
4». It is larger llian die egg of 1: nteholWNf Tonnw 
which was described and i Must rated bv Clltfic 
(W37). 

Second and third insLar larvae of most described 
species of h-t\\n\i>hhi have a more or less elaborate 
doisai shield tCumc IW). lo h. fluVHomi^ it 
consists of a sclcrolised modification to the second 
and third thoracic segments and first abdominal 
segment comprising a plate with sometimes three, 




("inii Wyig&l t£) 

I ig. \, Kelaiion.ship heiwocii total numhti ol cm^r^eiK-cs 
j..i 'jull fnf do t tixn\<'nnh< jta\ii tiivhi .ili <ne. and (fri / 
[lmi'On>\\ plus flt^Wtilletl I Fmcnnptcta v.tmix gyj| 
(Vcighl fin grains), reared between 5 March and M\ Apiil 
1995 from hiic,>!\i>m\ uinuthlnlrnsn nl GohlWtf Soiali 
Airslraki 

mosdy four d\-\\S rarely five anteriorly projecting 
prongs arising from the third thoracic segment The 
structure of the shield is similar to that depicted lor 
/-, hvkhit fit TonnOU" by CUfriC t 1*07), Its function is 
implicated In feeding and the enlargement o( gall 
Laviiies (see Discussion). Indeed many pellets ol' 
plain material were found ill the gall cavities often 
adhering lo the dorsal shield. Faecal matctiul within 
the gall cavities was nol observed. 

Emergence aiul Phetiolo&y of Fergnsonina and 
liwotittti'd Hymctiopicii! 

Gulls collected in late February 1095 contained 
only larval stages of h. fitivhvrriix but pupae were 
present soon after. Galls were caged in the field on 5 
March (day 0) and the first tidnlt / flavicniwfo had 
emerged by day 3. Adult flies continued lo cmetge 
lor about the nexi -Id days, peaking around cl,i> Is 
(23 March) (Fig. 4a). The 370 adull /. flavivtmus 
i hot emerged Irom the caged galls had a sex ratio 
slightly biased in favour ol males (1: 1.2). Further. 
ihc uinnber of flies that emerged had a positive 
reluliotiship with gall size (measured as dry gall 
weight) (Fig. 3. line bl. However. Ihc regression 
againsi gall size was steeper for Hies plus associated 
Hynienopiera (Fig- 3. comparing lines a and b), and 
the correlation slightly better (F = 0AM scimis 0.X7). 
compared with Hies alone. The reason lor this is not 
clear but, presumably, gall size is a reasonable 
estimate of available food and so for a given gall 
there must be a limit lo Ihc number ol I lies it can 
support. However, all jjalls surveyed (Table I ) 
contained some Hymenoptera, and these individuals, 
whether they are phytophagous, parasitic on E 
jUtvicomi;\. or both (inquillncs), will have displaced 
or killed flies that otherwise would have completed 
development and emerged. Hence, the steeper slope 
0/ the regression line for flics plus IWmenoplera 
represents what is a closer estimate tor die carrying 



134 



G. S. TAYLOR, A. D. AUSTIN & K. A. DAVIES 



:l Fergustmina flavicornis Mailoch 




0.5 -i 



e Euryioma spp. 




b Hmcon fcrgusotiinus sp. nov. 




0.8- 






t Megastigmus spp. 


0.6- 






0.4- 






0.2- 






0.0 < 




■ i • i ' i ' i > i ■ i 



C Ciirospilus sp. 



UJ 




0.5 ~i d Eiulents sr 

0.4 H 

0.3 

0.2- 

0.1 - 

0.0-cJhd-t 

10 20 30 40 50 60 




6 8 12 17 23 I 13 

March April 



30 
1 995 



0.5 ~| g BootiWL'Heus sp 
0.4 




T ■ 1 ■ 1 ■ 1 

10 20 30 40 50 60 



6 8 12 17 23 I 13 

March April 



30 
1995 



Fig. 4. Number of emergences per day for iergusoninu fhtv'uoruis (4a) and the six mosl abundant Hymenoplera (4b-4g) 
reared from galls between 5 March (day 0) and 30 April 1995 (day 56) on Eucalyptus camalduiensis at Goolwa, South 
Australia. The standard errors around the means are different tor each species because the number of galls in which Ihey 
occurred was also different (see Table I ). 



lilOl (KiV Ol- hhtiGUSOWNA AM) ITS ASSOCIATED HYMINOPTI K A 



155 



T Mil t I Total iiifti^currs (ruttt $ Xfan h SO April, IQ95 v\ Fcrgiisonina riavieornis and ax\in iahd hynH'ti<>i>unw spp 
from f-ergusoninu/i 'crjnisobia #alk on Eucalyptus cumul.lulensis uf fifltitwa, S. Ausf. 



Spo/lCS 

I'.'iyjiMwiiUi jl<ni, amis Mallocli 
Hrthon fi'(itns<>nintts sp. nov. 
hwiltoci vptU\ spp 
CnWiSpiitis sp, 
huU'nts JtJT. 
t'ctiii/hi>i.\ sp. 
I'mMoithi spp. 

I /'(/('( \/>(f //l\'/7/( f//( /,7 .AslllllL'inJ 

Dihnptnnh lit: sp 
Mi'vjisll^UiS spp. 
titnrfiincflciis sp, 
Tolal par.isilitHK 







3 


4 


Gall number 






1 


-1 


5 6 


i 


Total 


19 


2.4 
17 


!2 




44 




44 ISO 
1 10 


& 


HO 


T 


i 


31 


'1 


1 





1 


2 





r» 


(> 


7 


1 


1 


:i 37 


fj 


S^i 


i 


1 








X 15 


g 


44 








II 


1 


g )i 


fl 


1 


1 1 


II 


3 


I i 


4 25 


3 


71 


R 


5 


1 


i 


3 5 


5 


15 


U 








o 


4 3 


2 


9 


u 


s 


1 


19 


12 30 


] 


75 


1 





17 


2 \2 


U 


J3 


>3 


4-7 


7 


M 


5<> \m 


2K 


-VI 



capacity of u pall in terms ol lout! resources. 
compared with that for flics alone 

The composition of ihc hymeuopletan species dial 
emerged from caged galls varied in all eases, eu vpt 
between jails 5 and 6 which yielded Ihc same 
species {Tabic I ). (lulls contained an average "*' 
seven species (range s 4-9), wilh no gaJI containing 
all species (nole lhat in Table I species For each of 
/Wr//orrv/>//<v, fpnltnna iiitd Mexusltiimits have 
hceu combined). The biology uf the individual 
hynnmopleran species in 1liix guild is discussed 
below. Ol ihesc. several were reared in sufficient 
nunihers t> M) individuals} lo examine their pattern 
ul lancojeiice (l*ig. 4b-gi. Hancn fcn*ustmintih sp. 
nov.. Civmsptlus sp. and Mvftn.itlymm spp. all had a 
pKOJfc in their emergence prior to tbatof K.fl<i\*tcflnifa. 
Fui H, }cri>tn<mitm\, this puak occurred al ihc 
heginning ol the study (day 3} and virtual y nil 
individuals had emerged prior lo ihc peak \)\' F. 
lUivuornis on about day IS. while C/WMSpf/lM sp. 
and Mt#>asft$milS spp, eoiuimied lo emerge from 
Halls unlil day 56, alheil in low numbers for most of 
this time. /-.Wc/v/n sp. and BuryioWti N>P- bfld >' DtOftd 
range of cmeigcucc limes, beginning prior to the 
maximum in I. fU(\ i<*>nns (day I 8), peaking al about 
>\a\ 11 and couliuuing lo, or almost lo. day 56 when 
the study was terminated. In contrast, Bnotunrllfiis 
sp_ did not start emerging from galls unlil 21 days 
alter the peak in /.' fkiviconus. b'mergence ol" this 
species reached a maximum about day $$ and some 
individuals were still emerging from galls up to dav 
S(j. Ol |lu" oilier hyuieiioplcran species lhat were 
reared in Urn numbers <<30 individuals), I he 
emergence ol C mgivi'incta (days 1-27) straddled 
the peak for f; JlavUcrnis. fttecihtctvptu* spp. to 
individuals from days 23-39) and Pilropiitolt'fUt sp. 
i 1 * individuals from days 28-56) emerged well alter /' 
f/a\i<'t>rni\. while the single spedmen of ! s ttii<>hiu\ 



sp. emerged between davs 13 and IS 1 18-23 March). 

laxoiuimx tint! hiah^y uf Hyntcn"t>ura a^socian-d 
wiih I-ergusonilia 

During the study 12 species of Mvmenoptei'ii were 
reared from tTrxus<>ninti/fTrxtt\ohitt galls on /.' 
uitmtfifn/fu.sis This section describes some aspects 
ol the biology ot' Ihc species concerned and 
speculates on other aspects ol their assocuilion. 
based on information available for other related 
species. Notes are provided on their taxonomy, 
including I ho description i\i' a new species, and an 
illustrated key is provided lo ideniily Ihc 12 species. 

bamily Biaeoindae 
ftrmon fergitsoninus sp. nov 
' <ROS5-S> 

ilolofyi)v: V. Soulh Australia. Cioolwa, b-K.iii. 1995, 
G. Taylor & K. Davics. reared from Fi'r^ti\(*ninii 
galls on l:itt_ulyf>ni\ sp. (ANIC). 
f J tirui\>pc\: N ?V. IS 3$, same data a:- lutltUype 
except some with dates as follows 9- 1 2,ui.95. 13- 
17.iu.95, 24.ni -lav.95 <ANIC. WARD. 

Fetnola 

Si/e' Length 2.4-2.6 mm (excluding ovipositor). 
Colour Head, senium, pronolum, legs and anioriur 
two-thirds uf metasoma yellow-brown lo orange 
brown; aOteiuiae. eves, posterior parts of mesosoma 
and inetasc-ma. distal half ot hind legs, proximal, mid 
and hind coxa and ovipositor sheaths black; 
sometimes area around ocelli and posterior margin ot 
scutum darker, wings evenly inliiscaie. 
Head: In anterior view face half width ot head; lvi-s 
moderately bulbous, hairless: lace, gena and 
mandibles evenly covered wilh long hairs; in dorsal 
view vertex, dorsal temples and lateral frons with 



136 



G. S. TAYLOR. A. D. AUSTIN & K. A. DAVIES 




HIOUKil Ol nMilMtWNA l\ND US \SS<j( IMVMhHIl \OPT] RA 



i \y 



l&Ug hairs; in lateral view lower temple glabrous: 
antennae slightly luflgw than body: llngcllorociVMiH 
longer than wide, 

Mcsosoma. Stiit and scutcHum smooth, shmv 

with a lew long sparse hairs on posterior parts ol 
each termite, senium willi sparse hairs along lateral 
ltfU.rg.lH and illpjlg notauli; noianli indicated by hiTnl 
i1ept\ ■ssiou.s more prominent in posterioj half; 
seutcllai suture £tighlty depressed and curving 
posterioily. comprising 12 14 lovcalc puiMuivs: 
dotsal and lateral sculellum delimited by row ot 
loujqsh hairs. propodeum with medial longitudinal 
..i mm in posterior one-third, lainlly depressed in 
tii'L't'iH midline, with sparse long hairs in lateral 
purl in lateral view prouoUun aild mesupleiuon 
smooth and mostly hairless: cpicnLinial area covered 
with long hairs; inetapJeuron covered with long 
1 1. in-,, s hum. ih in dorsal half, faintly rugose in 
poskao-vcnlral part, lees evenly ami densely setose 
eu'vpi lor outer surfaces ol coxae, loic win;, wtlh 
IM and I Ks+Vl laindy bowed. 2-\is straight: '-CUI 
evenly curved: 2\w\ suhmarymal cell elongate, sides 
[^Rfl and 3-M| almost parallel 1st ciiseal cell 
moderately elongate 1 1 Rs+M 2,fl * length ..I'm cm 
Melanoma Mosily smooth throughout, sclcioliscd 
part of Tl slightly longer than wide, lateral margins 
with lung sparse hairs, posteromedial lobe ol II 
broad, family longitudinally striate along posterior 
margin, delimited by shallot lateral luiiows Which 
arc peicuirent to base of Tl: \2 with bioad rncdi.il 
longitudinal carina in anterior hall with a lew striae 
< ahcr side, postero-mcdinl area very faintly raised 
ami triangular. I? a\k\ J3 suhequal in length and 
comprising ahouL one-llhrd length of iiielasoum. w iih 
spaise long bans in laleml parts; TV i'5 aboul three- 
quarters length ol T2 T.\ with transverse row of Ion!} 
hdirS <md .a latcial margins; ovipositor and sheaths 
slightly longer thai) hods, sheaths with even covering 
of Icinji hairs, slighlls longer limn diameter ot 
sheaths. 

Mule 

pifl'erJDg from female as follows; Length 2 }2.(> 
mm; body generally narrower and moo: elongate 
pariicularly posterior mesosoma and Tl; scape and 
often proximal one-quarter of antennae brown, let's 
with same colour pattern hut yellower; wings 
nation l-i, 

('om/tw fii.s 

This is a very large genus in Auslrarui of which few 
Species have been described. Primarily, members ol 



the genus arc parasitic on lepidopieran larvae bill 
the) have also been recorded from Cok-optm. 
Oiplera and leal- mining pergid sawflies (QuickC 
I08S; Quicke & Ingram IW, AuMin & KttiliK 
l9S n i Other than one unsubstantiated record of a 
tlntton species being reared from a gall -forming 
homopteiail tChadwick ( *C Khkitin l*)7M, ihi.s is the 
first record of an Australian species heing reared 
from a plant pall. Studies on Hie biology ol numerous 
species overseas and from Australia indicate that all 
i. Ki. liter-, of the gCHUS are solitary primary 
ecloparasiloids. Given this, and thai H J'n\n\f>iiiiiti.\ 
was reared in moderately large numbers bom five of 
seven gaily during the projeti (Table I ). Ilus specie-, 
is most likely to he parasitic on fcwfjltxmtltu laivae 
PivmomsIv. ( mrie (KH7) rcponed tcatiug an 
unknown braconid from galL ot icri>ti\nt}itni m the 
( inheiia area, and this may he Ihe .same species as 
deenbed icrc Howevet there is .ippareatly no 
w.nt Ik * maie.-iat available From Carrie's study in the 
AN1C or elsesWieie. I nether, Cuirie . I'.'^7,i \hu\K\ 
this braconid "lo feed indiseriirhnalely "it gall tissues 
and fly larvae " We have been unable to conhnn 
t:uMic*> obseivaiious hut. il this species is a 
faeoltatise icc^r on gall tissue, il will be only die 
lourlh record t>l phytophacy 111 Hie iiraconidac in ihe 
world (Auslin $ Oangerfield niipttb. ). and the lust 
for the subfamily Braconmae 

ttnn*<m Jt'txitsiiHinits differs from other deseribed 
Australian $fttt*$H species in die form of the medial 
propodcum. shape of Tl. sculpturing on T2. length 
of the ovinositor. and its colour pattern. )i Jn caMly 
iLlculified from other parasitoids associated with 
l't^»ustmiiiit/t-L'v^Hs<>hitt galls by its complete wing 
venation i l-i^. 7). ciivnlar and depressed (cycloslomc) 
clypens (big. S). elongate Ov/SporflCjJ tlitu &) a mk\ 
cirong$ and black colour pattern. 

hamily lebueumomdae 

P(fr<ihnrxim{\ spp. 

ir'KiS^-lu 

't I lis genus belongs m an unusual suhf.umlv Hie 
Labeniuae. in that it represents one o! rhe most 
plesiomorphie groups of iehncumonids and has a 
Cfondwanan distribution tCJauld I0S3, 1 084; Gauld 
& Holloway I *->S0) /'c< ihu typtttt lias been 
postulated to be phytophagous on Ihe basis of the 
structure of Ihe mandible (Short l l >78) and il has 
previously been reared from gaWs on Aciuut and 
Eucalyptus (Gauld & Holloway l°Sb) and, more 
recently, from Ratiksiii (Auslin # Danjierlleld 



( ijis 5 s, lint, on (< n;uumrtm\ sp. nttv, . 5, 1)t>rsal hahittis (not showing ovipositon h. Silhonetic of dorsal body show- 
111)-' lenrlh of oviposiioi 7 l-ore ;in<l hind wings tvein r-m .imnved in hind wineJ, ft Anienoi vh-w of lieud sIkusii^ eii 
ml. ii cJypOllv Stale bars 0,S mm S, I »nni7: ()_S una K 



I3S 



O. S. TAYLOR, A. D. AUSTIN & K. A. DAVIRS 



unpub. ). However, only very few specimens have 
ever been reared, as is the case here (i.e. six 
specimens from four galls - Table 1 ), indicating that 
the genus is more likely to be a parasitoid or 
inc|iiiline lhan a primary gall-Former Surprisingly. 
two of the four known species of Poadoaypius, P. 
nignnnacttlatus Cameron and P. galliphflgus Gauld 



& Hollovvay. are here associated with FergMS- 
{>t}ifia/Feri>usobia galls. Both species have previously 
been collected from the eastern part of the continent, 
Queensland to Tasmania, but both arc recorded here 
from South Australia for the first time. Poecilocnplus 
nigronuuulatu.s has been reared from anthribid and 
chalcid galls on Acacia hngifot'ta and eriococeid 




Pigs 9-14. 9-11. Pocrilarryptits nigtomactttutits Cameron. 9. Dorsal habitus, 6 (extent of black markings indicated by 
slipplingh UK Silhouette of dorsal body of V showing length of ovipositor 1 1. hore and hind wings of 6 (vein 2rnc» 
arrowed in fore wing and vein r-m arrowed in hind wing). 12-14. Ciirospilus sp. 9. 12. Hind leg. 13, Fore wine. 14. 
Dorsal head ami mesosoma (extent of metallic green markings indicated by Stippling). Scale bars = 1.25 mm 9, 1 1:0,5 

mm 12-14. 



BIOLOGY OF FEROVSONINA AND ITS ASSOCIATED HYMENOPTERA 



139 




Figs 15-20. 15, 1ft, Emlents sp. V . 15. Dorsal habilus (pr = pronotum). 16. Fore wing (postmarginal vein arrowed). 17. 
20, Eurytoma sp. 17. Fore wing (sligmal vein arrowed). 20. Dorsal head and mesosoma (pr = pronotum). 18, 1°, 
I'ediobitis sp, 9. 18. Fore wing. 19. Dorsal head and mesosoma. Seale bar = 0.5 mm. 



i KI 



«. s lAYJ ni\ s i) AUSTIN &. K, A, DAVtl 5 



polls rui tM><thi>ttt\. white /■. snitlipM^as fins ix»ii 

associated wilh unknown galls on £ th'li-t>titc/i\i\ 
antl £ {Mm ///<"</ fGautd & Holioway 19X6). 
r<n( ihu rvputs spp. affi distinctive compared with 
OlHcP parashoids in the iniiM IjCCiUQjC Of ihch laiyc 

si/e lappomman-lv 10 excluding ovipositor) 

■and winy venation (* 7 *g, II). Although the fwil 
species arc very similar to each other they can he 
separated In the colour <»J the hind fcmui and l&rjgrh 
ill lite ovipositor 

1 amiiv Butdphidflc 

( 'hrosfti/iis sp. 
il-UiS 12-I4» 

Members of this very Urge ami laKonnmicrdly 
difficult cosmopolitan gcflllfi are relatively small in 
size and arc ohm hnyhily coloured whit metallic 
niatkings, as is the species heie Uilfetent species 
have been icaied as primary parasiloids ov a* 
lacultnlivc or obligate hyperparasiloids Irom leal 
mining and gJilMtomlns insects, Thumleri and 
Austin i I9k>4> avoided four species associated with 
ihe Mimes of I'lnhn Mtipkifict on cueulypts m South 
Australia which, in the case ol the donunanl species 
( ' muri>ttit\ii)in\ (jiraull. was shown tit be a 
lauillative hvperpaiasilotd ol Ihvnm and othei 
chalcidoids. The biology ol" the species here \$ 
unclear and it CGtUld he eithei a pi unary parasiloid on 
f <ytt\onimt larvae or facultatively hyperparasilic on 
one or more til lite gall associated Hymcnoptera. h 
was the most abundant parasitoid encountered dining 
Ihe study mid it was recorded Irom sill seven gfllls 
< Table I \, Ihe species is close to C \H ri()iii.\ Guaull 
bul differs in the coloui pattern of the dois.il 
mesosoma, and tints may tepreseul a new species 
With m the parasitoid guild associated wilh 
h'n;nvo»itni/h< rxHvnhia .ualls, it can he easily 
identified based on the form of the fore wing 
seiiiilini) < hiv- rJfc colour pattern and shape ol Ihe 

both (fig. |4K 

I tuLnts sp. 
I FIGS 15. m 

I. ike tVr/v'v/>/7//A, this is a large geflUR whose 
members are difficult to identify. Hxtralimilal hosl 
iccords indicate thai Undents are parasiioids of 
Icpidopteran larvae or arc hypcrpurnsitoiils. 
parlienhirlv of Uracomdac (Boucek 1988% The 
association of the species reared here Irom 
ri'tgnxtinitui/l-t'r%n.sobiti galls is ihe first biological 
information recorded for the genus in Australasia, 
GiVcB dial species overseas have been reared from 
Bracouulac, it is possible Ibat this species is 
h.vperparasihc tin &• fitffitwminiu This proposal is 



supported Ml least in part by lire tact that Jjtdrrns sp. 
was reared fiom galls that also yielded individuals |>l 
ft. frrL',n.\i>ninn\ (table I), and iha* no other 
hymenopferans m the same galls aie large enough m 
body si/c to provide sufficient food fur Ihe 
devclopmcnl ol Ltui.tH.s larvae iw'uh the cvcepliuu 
of rti<ith>ri\iHNy which was too tare), In "he 
parasitoid guild associated with FW'gMVfVli 
lin:u\nhiti >mII\, l.thlriu) sp. t -, easily separated 
Imhii oilier species by its metallic blue-green colour 
elongate body fFig, IS), nicsosoma with very Jute 
letieulate-punctaie sculpttnuu: and lore wine Willi 
posMoaicuul vein ahoui as |0Mj as \tiem J win 1 1 e 

141 

ft /Ht>hitt\ \/>, 
(FIGS IK. 19) 

Membcfs of tins feints are known n> be | afV 

and hyperparasiloids of eggs, pupae rind sometime-, 
larvae ol' various i usee is. mainlv I rpidoptcm 
t '.dcopicia, Ntpleia and Hymcnoptera (Boucek 
IVKNt. Several spettcs have been reared trOftl It I 
leediny insects on eucalvpts lc,e.. Cruhn lu^nn 
Walker ami fhylfU'W/ipho^tt J\Wf$f?tti Kiek - AtKti'i 
Ik Mien I9W: Thmnleil --V Austin 1'HM) ahd varnMis 
lealanining, Piplera ( Boucck l c >£8). bul apparenlly 
none has been reared from ^all-assm. iated host*,. The 
species here was teeorded by only a suiele speemuti 
(Tabic 1 1. Although Us biology is not known, iln*. 
very low abundance would indicate thai il is 
probably a hypctpaiasiloid on another hymenoplerari 
in the hi , ry,LiM>ninti/hfi\>tt\nhia gulh] Ptrftohms sp. 
can be easily scparatctl Inttrt the oilier parasitoi-.K 
recorded by its relatively ^huri ittnipact body H i 
IsU. blaeK colt at r. \ei> finely relieulale dorsal 
mcNosoma, and eloucaie posimarginal vein (Flg r I H> 

fanuly f.uryiouudac 
/ 'uryfnmu spp. 
HK1S I12TI) 

this is a veiy large yenus in Australia containing 
species with divergent hioloyies S^CCJC^ -an be 
phytophaguus. facultative!) parasitic, obli^atonlv 
parasitic ot hvpei parasitic (Botieek r>HS). In 
Australia, the niajoril> pi species are postulated to 
(all into the laltcr rwo ealCiiorics, bill the eeuetie 
.las' itiealion is not well understood and specie 
belonging Ui apparently ckisely related genera and/or 
spectes groups are known to be cither phytophagous 
or parasitic. A number is known to be eeloparasitie 
on ^all-foiiiunji; insects ku hyperparasitic l|ii(>uj. , h 
icIiueuiiHMiid and bracomd cocoons (Boncek I988J 
Currie { 1937) reported one species of Lmytnnm, II 
"vurirnjifrK" lo he phytoplui^iMis m Irr^usoniua 
.walls neat C auberra tvtirirnjipr\ )s an unpublishcxl. 



HUM (Mi 1 . t&toKtitWmM AMMTS ASSl/rt\lM)ir, YUSuHl.kA 



III 



manuscript name - sec Oahim- 19Kb. K«mCvk I9SK). 
I'nlnrKinutely- Wj! telVd not been able to evrtfifflfl the 
biology (if ihc- specif^ encountered in Iftis study by 
duvet itb^crvaiiim. One of Lhem twee below i nui\ bo 
I he same tocclcg (\j lecorded by Curric (N,B. no 
Vquohei material is available bom hi\ -attdy) Jiitl 
therefore possibly be phytophagous (see Piscussinrrc 
01 they could be different speeies ttnd be parasihc on 
h r^i\nninn larvae or on one of the othei hymenopieoins 
.n the guild, possibly 8. fei*giUWIlJtni$ sp nov. Th: 
speeies here was recorded fmm all (he calls surveyed 
and was the third mosi abundant hymcrioplcran hi 
Iht' h'i't'ifHSDiiitui/lnyjisoMa guild liable 1 k 

kttrvt"nnt often display iulraspecific morpho- 
logical variabilis and HfC therefore diiVieuh to 
scpai. ae al Ihe speeies level, The material reared heie 
varies slightly in st/e. eolour. pilosiiy and seulplu in: 1 , 
and may represent more lhan one speeies. I he genus 
can be easilv separated from other parasiroids in the 
guVy based on Ihe shape <tf ibt- pronuium, dorsal 
sculpturing paiiern 1 1 ip 2i)), and wing \enatimt | Fig, 
17 \ The sexes are highly dimorphie wilh males 
heme smaller, having a dispropoiiiunutcly smaller 
mulusoma, and asymmetrical pilose antennae, 

bamilv Plcromalidae 

Ci'i-huvhtt tti'jr<H incut Asbmead 
|HIGS23 ( :4. 

This speeies and all members ol' the genus i,\rc 
probably inqilflines of hynienoplcrun and dipleran 
gall -formers on AtH&titl and i.ttiUl\}>Uis HJouccW 
IVH&l. Previously. C. tiitjitu iiu'ta has been reared 
from Ihe galls ol Ihe phytophagous ptctonUuN 
/Wr/l/frftA&N'f Hfld Ffifflfimptfta (Noble Wk I'M I I 
and. m ihe laiier ease, has been proved to be an 
iiK|uilme. i.e.. il kills ihe resident ceeidogenie wasp 
ami forms its own eelK to teed on ihe gall tissue The 
species was originally described fi'oiu material 
reared from ^agromy/id galls on fUt<rtiffp{tti 
i<>fvmbo\<r (idemified by/ Boueck f 19891 as 
h^nsiuiina gallsi in Sydney (Ashincad 1 91K) 1 . uixl 
was %ubscc|uciltly recorded from fu'tyjisutthm/ 
/<\i<-n\<>hitt gaits as. an obligate para%itoid in die 
Canberra area by Cuuie H937). Specimens in ANIC 
were collected from galls oO £ V&M&ldtiwlwh »U 
Alice Springs, NT antl fc va7?en L. Johnson and A. 
nmyyuhtiiitu Lahilk at Biehenn. Ttw. (1. D. Nanmann 
pers comm.) Wc have been tillable to confirm ils 
biology in this. sludy T but clearly il is mosl likely to 
he an im|uihilc or primary parasitoid ol /. ryjtwiiftii 
O't'lthxhti ntyrccitwut was reared only in moderate 
nnmhers he-re but il was recorded from live of the 
seven galls smveyed (Table I), Il is apparently 
distribuied along the east eoasi of Australia from 
north Queensland to Tasmania, but litis seems to be 
Ihe firvi record of the species Irom South Australia. 



Oji'lttcyivt m^tvcu'i (a Bin he obftily -cparaleU lr 

oilier parasiloub associated uub I tfytt\tnntni/ 
Icryttsofrnt trills hv its elongate sm:maj \em. shun 
oviposilor and disli.utae bn.el'it yellow bodv and 
tuelasonia wilh bhuk InOintdW! >lripes it-cj 1 ., 

DitmpiiHtwIfd sp, 

I PICK 3b| 

Members of tHi<< tienus liavc prc'i"u,^|\ Iveu 
nvuicd under ihe Ibiyimd.-te rti as misco\:a.si. i : 
|i|eii>maluls. bin mosi recently they have been pUi |cd 
in Iheii own suhfaniily. DhtopJnOWJ'iniW wiihin the 
IVMumalidae iRouC'ek I'>88), lYeYiou-.ly, ihcv HftVi 
been reared horn \t>ioint'}j>Jni giflls ^m /ja'alxplits 
<\\\i\ unknown ^.ills on / '/c<</v/'/rv, AiUtia, CaMtunun 
(tiouc-ek l'>XS) umI a number ol othct plains. The 
bioloyy o( Ihe tieuus is "'Oily kii-»\Mi HuUrcia 
fume i IVS7) reported /.» t utttytyMvatitm (>unov t\ 
be phylopha^tms within i<:ryn\ntiiiiti/it'tyuxt*htu 
>:alls in ihe Canberra area. During diis study, ouls 
nine indi\ iduals were reared from three ^alls (Tnhlc 
1 1 tfrtdi based \^ Ihese numbers, lilt Spficlc^ hi moiv 
likely lo he tin itiquilitic nr parasite on \uie oi -moa 
gilll inhabilanls. We have llpl been able poshivel> 
to itienify the species heif as /). mmfitvwiyiitiris 
primarily \.\u^ I" moiphulooical variabiliiy amonj. 
species in the genus (Boutck l^l^i. Howvvet; it 
can be easily distinguished from Ihe othei parasiloids 
in the guild by us brigfK metallic peen body 
lalerall) e< impressed melasoma. and e\scrted 
oviposilot (l ; ie. 2M. 

harmlv b.iymid.h 
,W('e,//.\7/i > ( y;u/.s spp, 
.HCS ^1,2125. 

The Mc^asUgminuc aie best characterised by Iheii 
laree. knob-like foie WUIg stigma and elonealf 
Qvjhnsiior ol the female. The genus McyuMi^/nus is 
laaie itifttl ta,\oiM»mically difllcult in Ausiraliu. with 
more lhan At) described speeicv As well, its members 
have diserse biologies m thai apparently closely 
related species can be phytophagous, ceeido^enic. 
inqmlines or ]>aiasitic- althoueh the detailed biolotyv 
of no Australian species has been confirmed ( Boucck 
| l >x<X), Various species have been reared from dipleran 
ealls mi A\l<rm'nnh>Mt\, Aftii»Hi>tplui iLriocoecidac) 
on Eitailxptus, Bnuhophaya.\ (lui\tumidae) galls oh 
Ircmni'tus, i'ntlnloitiiMcr iPteromahdaei galls mi 
Ai-acia. as well as unknown galls on numerous native 
plants (Boueek J 988k Tw<> species, \t tjulnifuase'tut 
Oirault and Ahyti.siiynius sp.. were reared from 
h-ryu\('wtm/J : t:iy,H\ifhni galls in the Canberra area 
by Currie 1 1 L )M I, and were reported by him to be 
phytophagous. We have been unable to confirm 
Cume's identification <>l these species, or to show 



142 



a S, TAYI OR, A. D. AUSTIN & K. A. DAV1ES 




Figs 21-27. 21, 22, 25, MegdStigmus sp. 9 . 21. Dorsal head and mesosoma (pr= pronotum). 22. Fore wing. 25. Lateral view 
of metasoma and ovipositor. 23, 24. Coelocybu nigmcinctu Ashmead 9 . 23. Lateral view of metasoma (extent of black 
markings indicated by stippling). 24. Hind leg. 26. Durop'moteUa sp.9. Lateral view of metasoma and ovipositor. 27. 
BaQtanellus sp. 9 . Anterior view of head. Scale bars= 0.5 mm 21-26: 0.25 mm 27. 



mOI.(Ki\ Ol }UV>(<Vh\lXA:\\n 

that ihcv are indeed phytophagous. The number ol 
individuals obtained for (lit* two .species reared here 
whs pooled because of initial prohtetQM in separating 
litem ia.\onomteally. Combined, they were reared 
from all seven galls surveyed and occurred in large 
numbers fluhle 1 |. being the second MOM ahundant 
genus present in the guild. 

Mct><i.\tiXmit\ ean be easily distinguished from the 
other lt j txn.wtiin(t/Ti'rga\obia gull inhabiting Hymen- 
opicra by their spot-like lore wing stigma (Fig, 21\. 
bright yellow colour with some dark markings, **ikI 
the length of the ovipositor (Fig. 25), From each trite 
ihey ean he separated by the eoloui of the metnsonui. 
The metasoma of one species i.% mostly dark vvheieas 
111 the OthCf fi '* VtJl<W With transverse black stripes. 

KtfOtUtu'lh'HS Sp. 

.FIG. 27) 

This is a moderate-sued genus Of inegastigihincs 
characterised by the presence of a medial tooth on 
the lower clypeai margin and some metallic 
coloration. There are 20 described species in 
Australia ol' which three have been previously 
associated with seeds ol Cusnurin>A and gulls ">u 
< i;ni\ and Munu tints (Boucek 1988). The biology 
ul the species leaied here from Ftrgiistwhut/ 
liivitKi'hiti galls is unknown, but it was reared in 
moderate numbers 1'rom three of the seven galls 
e\ammcd (Tabic I f, Based on these numbers and the 
I.U'I Iri.u it emerged well after all other call 
inhabitants, ihis species is most probably a parasm >id 
or iin|udinc- The species is very striking and easy to 
separate front the other Hynienon-rcrn associated with 
I rrxtisonitHt/rcixHSithia galls based on its yellow 
hod\ with bright metallic green dorsal head and 
mesosoma. black dorsal nietasoma and clypeus with 
.i media! tooth ilig. 27l 

Key to the pai asitmds reared from 

F'frKUSfmhi(f/l ; 4'rt>nuthia ( oills 

in South Australia 

1. loie wing with more than one enclosed cr II., 
moderalely complete venation (Figs 7. 1 I r. 

prepeelus absent (Ichneumonoidea) 2 

FOtfC wing with m more than one enclosed uifll. 
venation restricted to anterior margin ol wing 
it igs H, lb. 22): prepectus present 
tl'luilctdojdca) , 4 

2. Tore wing Willi \em 2mcu present; hind wwg 
with vein r-in meeting Rs after Ks diverges from 
Sl+R tPig. I I ) (Ichneumonidae) | length about 10 
mm without ovipositor; ovipositor shorter than 
metasoma (Fig, KM: clypeus normal; body 
orange- yellow with black stripes on metasoma 
and bind legs (lig- ^)| (/V; f v/7<M n/jf//\/ 3 



ITS VSSOCIATb.l) 1 1> MKNOPTRRA I4S 

Fore wing wilh \ein2mcu absent, hind wmg with 
vein i in meeting Rs before Rs diverges from 
Sc+R (Fie. 7) (Braeonidae) (length about 2.5 mm 
without ovipositor: ovipositor almosl as long as 
body Fig. bi; clypeus areola! and depressed 
(Fig. Sf; orange in colour with posterior 

Uiesosoiiw and metasoma black] 

,... ..Branntji'iii>n\t)ninus sp. hoy. 

3. Hind coxa black; ovipositor about 2,5 x length of 

hind tibia , /' tn^romrunlatus Omenin 

Kind coxa yellow: ovipositor about 2,2 x length 

of hind tibia 

,,./' xnlliphtityts Oauld & llollowav 

4 In dorsal view, pronoturn large and sub- 
rectangular or broadly munded (I igs 20. 21 ) ,. ,5 
In dorsal view, prouotum narrow and composing 
littles if anv of dorsal surface of mesosoma (Figs 
15. 19).....'. ".% 

5- Fore wiug with elongate stignial vein (Fig. 17l 
(Hurylomidaet |body all black, Ices often brown 
or orunge-brtiwu distally; female with dorsal 
mesosoma coarsejy rettculale-punclatc (Fig. 20). 
ovipositor short and virtually hidden under 

metasoma) Lurxwma Spp, 

Fore wiug with large spot-like stigmal vein iiiiv 
22); female with elongate exposed ovipositor 
(Fig. 25) (Torymid-'ic) ..- -..- - 6 

0. Body all yellow with some dark markings; 
clypeus Wtthotlt a medial tooth (<V//-v</s//.e/w/\> 

Venlral antl lateral surfaces of body yellow, dorsal 
head and tnetasonia bright metallic given, dorsal 
metasoma black: clypeus with medial tooth (Fig. 
27) i7t y Rotttdtielti'US sp. 

7- Metasoma mostly dark dorsally. stripes or striped 

pattern indistinct $t?$it$ti#l}(iJA sp. i 

Metasoina yellow with rransvetse black swipes 
MestislisiitiUM sp. 2 

N. Hind tarsi 5-seemenlcd (Fig. 24) (Pleromalidael 
" _._ _._ _ 9 

s*. Hind tarsi 4-^egmenled fFig- 12) (Fuiophiilaei 
10 

9- Body bright yellow or yellowTirowu. melasoma 
depressed, with bioad black transverse stripes 
dot sally, ovipositor short, hardly protruding from 

posterior metasoma (Fig. 2\) , 

Ci'tfa vhu Hixrochti to Ashmead 

Body bright metallic green: metasoma 
compressed; oviposthM e\ sorted posteriorly 
i Fig. 26» ,. Diir<>pitv>lellu sp. 

10. Body with distinct bright eolom pattern, 
including metallic green markings oo dorsal 
mesosoma (Fig. 14 1 and black transverse stripes 

on metasoma - - Cirwspihts sp 

Bodv with uniform dark colour I t 



44 



■ I W ••! U1AI ^tlNA-K VhAVIlS 



ii, body ciouguie irg. 15) iiki.iIiil hltit LTtxit in 
, olour dorsal mesnsoma Willi very line 
iviu.ulate-punuuic sculpturing; ftufi wing Willi 
niwMTttrgtnal vein ntmut us lt)tfj as sijginu) \ -<n 

'I''. 1 - 1 1-6) l.tnUiH- iff 

(jivlj iclalivcly shod iMtirpaci I Fie, 19) I 

M.uk in colour; dors..i nu sosunvi with vciv ITnw 
icncelaic sculpturing: lore vviiii' with 
■ .turginal vein nrOdi l.nKcr dun ^Igftihl VOID 
(Frg. Nil r>.;li<>htns sp 

Discussion 

TV* mechanism:, by which ififc Imji ptoni * < Ite aiy 

ni.in-.-cd do profit tirult Lu form a naM remain 
problematic PuiTie ( WJ71 -obwrV«] ihitl nematodes 
■\k\>: aciivels lirdillg jiuI cauviiit; cell proliferation 
ol plant tissue around the ll'-js.'MK fly before they 

'i.i | [Wit, I Ills soggesj'. l|l,|( itt^itllr liriilHliHlcs [fl I 

nwponsible lt)i gull initiation (presumably via 

,.ilK.il\ seueiions). ll is llO( UkOUccivahlc llu\ve\s . 

thai ii I ruction of ovipositiott lluid whiJi is injfii i 
imo the plant by the Ily along with ihe cgg\, (V 

ii " <. IfLMML'Ml dalliaec t<> [lie tWIpOMtor alOllC. at ^ ♦ 

causes cell pi oh iv ration, li litis teen Libwsrvetl wllh 

oilier tvriitisotiiti/i/t irxttsehiti associations iDavics 
impijh. i thai gu)(S in which Ily larvae have died bill 
in which nematodes arc still presun. .vouir | n i I 
and pi able. Cumc ( t'H?) reported lhal when mlcrhlc 
Ily eggs were dcposjled, galls were inhaled, hm weiv 

'ahoited s( an early stage while ihe nematodes 
persisted foi some monihs bill cvcninally died HU1 
He suggested iluM rurlhcr gj|] glOWlh ffftf 
aUi (billable lo Ily larvae. Whalever Ihe mechanism 
-all L'lowlh lor /-.' Jht\'ivotvis was particularly rapid 
ill thai galls attained lull size in a mauei of wcek- 
dming (he late summci ecncrulioti. 

While n<> lacval material was observed in gal] 
Cavities Of /.' flnvu onus, (urric < I *->37 > ohserved lh.il 
faeces were "practically absent" In 'ill hul pup species 
111 / i r.:tt\i>itituL Me postulated lhal nematodes, which 
hy then occur within the llv cavity, teed on llv faeces 
llius keeping the cavity Tree of waste. Given the strong 
stylci of die ncinarodcs. it is more likely that they Feed 
.'ii plant cells Microscopic examination of specimens 
eleaicd m glycerol indicated the presence o! a retluni 
and anus in h. Jhivic<>rf}is larvae. It is possible, 
however, that larvae may store, of produce minimal 

mounts of euiciuiy pnxluets to avoid fouling the 
Cavity. II ibis is so. u suggests an adaptation similar to 
that found (ii lite gall-forming Cecidomyiidac 
(Dipteral, the Urvae ft' at least some t>t wliuti pi^srss 
a FunClloMi ^*M"> bUI pfOdllce no intlieeable Avaslc 
pioducis (Gagne l u SSM. Perhaps related lo Ihe storage 
of wask h'ri>it\rttiit)ti ptiparia were obsenvd anchored 
within the gull cavity In means of a gelatinous 
substance ai the posierioi end. h is conceivable lhal lhi$ 



substance tepieseuls digestive waste viml^d just prim 
to piipaiiou taihke sonic cecidomviul ^-alls o iai'uc 
I'JK'M. fungal colonisation o! fresh t'Cn;u\<»iinti galls 
was noi L ( bscArif Ihe f|y larvae niav use llic dot 
shield, wild its anlcnoily directed prongs, lo scrapv llu 
observed pellets of plain material from the wall of ihr 
cavitv C imtic i \*>M i described third insiat Oy larvae as 
"iClUiuj: down" inner lavers ol the cavities for forttl 

I ho -Jiald iiKiy uJSO BO used (oaiidioi the taivae ui (JlflC 
spot within Ihe cavity and lo break down parts of id- 
cavity wallspiiortopiipalii»n t.ifaeiliune emcigeiuv >4 
theaduh Ily. 

ll is unclear how adult Hies escape from ihe call,-, 
after cineignic (torn pupana. Pisstctk'h ol 'jall- 
re\'eal*:d iluu bv ihe lime ol pupation, eavilies on Hie 
L'dgEs of [\\c \Hi\\ \vere scpaiatcd Inmi the outside 
only by a thin layer of epidermal cells, suggesi". y 

Mi .1 ll.c IflrVae lUlUirl nr.«i fc| tllO -Ulf-ii' |'n«.r lo 

pupati'Mi. Hresumablv ttUilJI Hies arc able [o ijjjc 
Ihruueh Ihis Hun layer relaii\cly easd;,. Lim ,■, 
deeper in die mature call ;irc scparaicd b\ ihm laycts 
of paicnchy;iimon> n.^uc nnd WWfflinxfi fmen tuii> 
each oiher. 

1 7 hc 12 sjtccics (»l HytHCii-'pleiM uared Mom lt*al 
galls of F, jhtvuorms during Ihis study DTprcwni I v 
ihe iiumlvr presuuisly recorded from Irixustminu 
ealls. Cunie (1937) reared imly si\ species frwm 
lliuwi galls in the t anbena a<ea. while Hams ( I^S. 1 ) 
recorded foui species from Ftir&WiOtmtl galls m InUi.i 
Further, several species aie here associated Willi 
bvrgustHHiHti grills for the h~rst tunc, \\/... B<m Hindi t'ti.\ 
sp., Cinospiltis -.p., h(//A'/v/v sp_, Prtlinhiit\ sp.. ini 
tWO species i>t r>><>>. Hat r\('lu\ i/' tiixr<>htii< itiaius and 
/' iHtlliphuiWs). AILhough ihe composilion oi 
hymenopieraii spci tB5 associated with indi\itiual gallH 
clearly dilTeis. the most common species encounteicd 
in ihis sludy, linosp/h^ sp . /u<rvti>tn<t spp, and 
Mcxastt'wtuts Spp*„ were reared Irom all gulls (Table 

I I while 13. fti¥$itMtftimnti <■' mxroi tttt ut and 
/ utletm st>. Were reared from more than li)% u\ 
^alls The <lillerence in die M^e ol the hymcnopiciau 
etiild between this study and lhal of Cunie i I9.s7» 
may be aitribulable io differences in ihe suuciurc aad 
position ut jjalls, i.e.. p ltgvfctiWU4 fttfllW leaf ,ea)l-. 
while Ihe species sluthcd by Curric formed bud gall> 
Also, the fact thai galls were puged in ihe field during 
this study means ihat all Mymenojitera were 
yollecieif whether lllcy were common (jr r t uv. To 
assess the composilion of wasp species by dissecting 
<jalls. as apparenlly undertaken by ( "'urric for at 
leasl pufl "I bts studs, is hkcly lo lead to an undei- 
esinnalionolThe true uumbei present. Such dissections 
ni'>slh ivveal larval stages lhal are often dillicull 
tti identify: they arc sofl-bodietl ami therefore are 
easily damaged and iheie is a greater chance ol 
overlooking rare species. 

The leaf galls caused by I- fht\h OPtlU llpOt R] • 



rioUKVi oi tmOU&OMNA and its AssouATPP HYM&NOPThkA 



115 



djBsccUon, to have a relatively toonTugenpnus matrix 
and. ailhoUgll SomC woody (issue is apparent, iho 
most imporlanl mortal ily agent is likely 10 he 
parasitic wasps, given thai as many individuals of 
I lymenoptcra as k ftuvtvomk were reared from galls 
(although not all species arc nceessaiilv parasitic 
see above >. Ilowevci. GUttlc I \ l ffl) attributed up lo 

6irv of Fer#mmitm mortality m tad gulls W the 
inability ul" fly larvae, prior hi pupation. t<> tunnel 
through the woody gall tissue induced l\v 
liymeiiopleran inqui lines, resulting Ml their 
Liilomhnieul within lite gall. He reported ihai wasp 
imp u lines were found mainly within (he OUier 
suilaee of hud galls, hut Hy larvae deeper in the gall 
WCW largely (wc ol parasites. He also repon d ilkit 
approximately the same number of wasps and Hies 
emerged from leal galls on & Sltmrthtmt KvVk 
islet, and Miggcsled thai there is a higher level ot 
parasitism in stem tip and leaf lip gulls than m flower 

hud galls. 

It is clear Iroiil this study t uu\ as reported elsewhere 
feg Houcck I *>8S; Cough & McMahon [9Sft Unit it 
ts very dilTiclill ro determine ihe biology ol" gull* 
assoeiated Hymenoplcra. Primarily, ihis is heeause 
then pie-adult stages are hidden within the gaH. then 
biology may change from <.mw siage to another las in 
TllC ease of iiiquiliues which may kill the resident 
gall-former and ihcn beeome phytophagous}, or it 
may be different Irom ivlated spceies Ihat do not 
inliahil galls. It is therefore naive 10 assume 
hioloutcal trails far ^all-associated wasps by 



cxtiapolaling Irom other species or inch reel 
information such as emergence lime and/or number 
ol individual present, Por such species, their 
biology may be determined only by direct and 
detailed observations including then morphology 
arid life history. In llns respect, we have been unable 
lo confirm the biology ot any species icai'ed in this 
study, although wc speculuie above on vvhul some ol 
these species may be doing, as ;i means ol 
stimulating further study. The comments made by 
CuiTie (IV.V7) that Ihe species ol Uifropinotrlhi. 
McfiHIfifittHW and Euryhnna he recorded from 
l\i-nsoi}'tiui bud galls are phytophagous musi be 
treated wilh some caution. given that he presents 
little ov no evidence lo ftppOH this conclusion, and 
at least Ihe latter I wo genera arc known 10 contain 
species which are cither phytophagous or parasitic. 
Purlher. his reporting thai f. nixivicuntct is a ''true 
parasite" and that the unknown bracomd tpiohabtv 
lUiuon) he reared was an uiquilinc is inconsistenl 
with previous studies on Ihese laxa (sec al>ove). 

Acknowled foments 

We wish to thank John LaSalle, r.rie Clrisscll. and 
David Wahl for assistance wilh identifying some of 
the parasiioid .species. Peter Cranston for identifying 
I. iftiviccrtiis, Peter Kolesik lor useful discussions 
and Paul Haneerlield foi ihe line drawings. We thank 
Lee tlallet for RCCdKS io !hc Held site. 



References 



AsnMt so. \V II. I 1'MHH ftflTW *'ii MWW N r ew Zealand it- 
Aienaliati pitrwiffc llvmcmtptrra. wilh desciipiioit- -'I 
m, \\ rnri:! ;unl new S^tft »es fifth', tjfUl So.', \ K W- <»6. 

I7K jon 

At Mis. A, D. tk Ai I in. O. R ( PiftOj Pam.Ml.nds el / Whti 
UtgSfl.i Walker (l cpidupinn: Nocluidaei III South 
AuMi.ili.i. wilh description (>] lW"i AC* SpOOle* ftl 
KnieoimU*. ium\ A'. S<><. V Au\i, 113. 169-184 

& Ku'ii's, W- t P>s l) » 1>vii new AuslraRjiii 

species vs\ OfttCtm ( (llymcin)pleru liou ■otmlaei 
pin;is(hc tin I'hvhu h't>pjni t i;it spp, ( 1 1> mciiDplcrj 
IVrpiiUw ). 7 Aasi-mt-Sm IN, 2117-21% 
liou t K. /.. ( P)NS)"Auslr.(linn nnalL-idoitJcii 1. 1 ly meeiop(cra). 

a Ku'vy^U'inuiic RuvwitwnJ'OmicmoJ Htaatccn himilies, 

with a Keclussiikalion ol S|iceies" (CAP hiiv.-muiioii.il 
Wallin.elouh. 

tiiMAvi'h. C I- K NiKiir-j. m. I. (f^76) Records ol 
parasilisin in the lamiln-s UhcmiMMitdite, Rrau»mdac und 
Autaeidue (I lynicnopicnO. ./. AliM rtlt &*•« l >- -X-M. 



t I 'imn C, A 1 1037) Oiills i»n Luailvpltn ins'-. \ ||p\v i>Tv 
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S«< V, IW. 62. 147 174. 

I'lvxns K. A.. & I JWO, J MiWfi) Nem:iHidt:\ jMHUldAill 
wilh Dipk-iii in South Australia U -if\s Hpecics ol 
hrvjtsiihiu Ciimc lutin ,i kiyusoiiinid and a new letiUn-l 
Ml Ssvfthflnmtt LuunTunrJ & Uyira ftoin u syrphui. Tntw 
K. .Sor. \. ,\a,W, 1211. I Via 

t)\n\is T. f IIV.S(0 A checklist ttf lypc-s ul Aitslrahiiii 
Hynien^pL'ia described hy .Mc\Lnio*re Arscne * nraull; IV, 
Chaleitloidea specie-. N ■-/ limcJ ^hci'j With ad\ 'isor> 
tioics pltij advlcfida and coitijjendit Mrm Ohl Mas, 22, 
;|o 7Vi 

FifalBfti -f M. & Nn>.i I. W R i PJ6SH)n ttW L'laSUafiejIltall 
Olid life histoiy ul liixnsul>iii , uniri iS[iti ( teriil-ariKl.ic: 
r^-nl,iluda|, i'nn, fh'hnitlth"f. S". . \\n\h 35 41) 4d. 

(i\oM, R.I (PiHyi I he plain feeding pull nudges til 
North America' it'uincll tini\crsiiv Pies 1 -. Ithaca. New 
York i, 



146 



G. S. TAYI..OR, A, D- AUSTIN & K. A. DAVIES 



Gauld. I. D. ( 19X3} Classifiealion. evolution ami 
distribution of ihc Labeninae, an ancient southern group 
of Iehneumonidae (Hymenoptera). Syst Enf. 8. 167-178. 

( 1984) "An Introduction to the Iehneumonidae ot 

Australia" (British Museum (Natural History), London). 

& Hoi ton. B. (1988) "The Hymenoptera" 

(British Museum (Natural History) London and Oxford 
University Press, Oxford). 

& Hoi.loway, G. A. < 1986) Australian 

ichneumonids of the tribes Labenini and Poeeilocryptini. 
Bull Krit. Mas. Nat. His!. Enl. 5$, 107-149. 

Goioh. N. & McMaiion, R .1. (1988) The biology and 
control of gall forming wasps on Gernldton wax pp. 1-6. 
In "The Production and Marketing of Australian Flora" 
{The Western Australia Department of Agriculture, 
Perth). 

Harris. K. M. (1982) First record ot Fergusonimdac 
(Diptcra: Sehi/ophora) outside Australia: a new .species 
of Frrgttsonhta on Sizy$ktw in India. Sv.si, Enl, 7, 21 l- 
216. 

Mai torn. J. R. 1 1925) Notes on Australian Diptera. No. vi. 

Enn. Linn- Sac. ftJ&W, 50. 80-97. 
Nalmann. L D. (1991) Hymenoptera pp. D16 1000 In 

Naumantt. 1. D.. Came, P.B., Lawrence. .I.F.. Nielsen. L. 

S., Spradbery. J. P.. Taylor, R. W., Whittcn, M. J. A. 

Lilllcfohn, M. J. (Eds) "The Insects of Australia" 2nd 

nxU), (Melbourne University Press. Melbourne). 



North, N. S. (1940) Tridiiloxasier acuiiac-lan^ifaHae 
(Froggall) (Hymenopt.. Chalcidoidea), a wasp causing 
galling of the llower buds oi' Acacia longijoiia Willd.. A. 
finribunda Sieber and A sophtnw R.Br. Trans. R. eta. 
Sac. Load. 90, 13-38. 

__ ( 1 94 1 ) Jrichil(>,i>aslcr nuiidcni f Froggati ) 

(Hymenopt., Chalcidoidea), a wasp causing galling of 
the (lower buds of Acacia implcxa Benth., and A. 
maideni K.v.M., with observations on Australian 
chalcidoid wasps. Proc. Una. Sac. N.SAV. 6ft. 178-200. 

QuickC. D. L. J. ( 1988) Hosi relationships in ihc Braconinae 
(Hymenoptera: Braconidae) - How little we know. Ent. 
Sac. QUI News Bali 16. 85-92. 

___^ & Incram, S. N. (1993) Braconine wasps oi 



Australia. Mem. QUI Mas. 33. 299-336. 

Schonrogoe, K., Stonb, G. N. & Crawtfy. M. J. (1996) 
Alien herbivores and native parasitoids: rapid 
developments and struclure of the parasiloid and 
inquiline complex in an invading gall wasp Andricus 
qiwiritscalicis (I Ivmenoptera: Cynipidae). Ecol. Enl 21. 
71 -SO. 

Short, J. R. T. ( 1 978) The final larval inslars of Iehneumonidae. 
Mem, Amen Enf, Inst, 25. I -508. 

Tiiumitrt. T A & Austin. A. D. (1994) Biology o\ 
Pliylacteophaxa frawatti Riek (Hymenoptera: 
Pcrgidae). and its parasitoids in South Australia. Tnm.\, 
R. Sac. S. Aits*. 118,99-113. 



STUDIES ON THE SOIL-INHABITING TARDIGRADE, 

MACROBIOTUS CF. PSEUDOHUFELANDI, 

FROM SOUTH AUSTRALIA 



By Alan F. Bird* 



Summary 

Bird, A. F. (1996) Studies on the soil-inhabiting tardigrade, Macrobiotus cf. 
Pseudohufelandi, from South Australia. Trans. R. Soc. S. Aust. 120(4), 147-154, 29 
November, 1996. 

A tardigrade isolated from agricultural soils at Avon is the first member of this 
phylum to be described from South Australia. Specimens were isolated from freshly- 
collected soils and from soil that had been stored dry for three years. 
Live and fixed specimens were examined under the light microscope and fixed, 
stained and gold-coated specimens were examined using the scanning electron 
microscope. 

Key Words: Macrobiotus cf. pseudohufelandi, anhydrobiosis, microscopy, 
tardigrades, birefringence, biocontrol. 



Itwtsortums ttf ihr AW// Stnirt\ <t( V \it\i | 1 l »o u i, 120(4): 14/151 

STUDIES ON I HE SOU .-INHABITING TARDIGRADE, MACHOHIOIVS 
CF. PSEVDOHUFHIANDL FROM SOUTH AUSTRALIA 



bv Ai.as K EHUD-? 



Summary 

BiK'n. A.I. 1 1 9961 SUiOics pfl Ihc sotl-inhabitme luKligriKie. \f,uiuhi»tt<\ i_f ptmud&ftufHttirttl* 'ruin South 
AustBilm, fnttts. K. Sot, S. Am. 120 (4), 1 47-1 H - 1 ' November. Wo. 

A lurtLigrade isolated from uiriculuirul soils ;ii Avon is tliL- first member of thi.s phylum to be- described from 
South Australia. Specimens were isolated from heshlv-collectcd kotft ami Ifcxn ftOal thai had been stoied itiy lot 
dim years. 

lue and fised specimens ueie evauuncd under the Ihjht imuoscope and lived, viained lu\k\ tiold-eoaied 
specimens were examined U&illV the scanning electron microscope . 

I his tardigrade, a stout cylindrical organism about 5(ft) um k>rtg by 1 51) uin wtde with the loin pairs ol stuhby 
legs ending in panvd claw*, lias been assigned to the species ihunthhuu* Cjf. ffm mioftiiff/tnitii on lite basis ol' 
the morphology of the bueeopharnyccal apparatus, claw shapes ami Cgfi processes. The stylets .ire slightly 
curved, sabre shapcit structures about 40 um in length and exhibit marked birefringence under polnrwd light. 
When these tardigiades are killed Lite stylets break down and disappear 

A specimen i>\' \f, el. i>,sauf<>ltnitittmli was observed feeding nn a neural ode and a Kignibcanf decrease in the 
number of nematodes in the sod as tlu- number ol unl wades increases has been demonstrated. 



bllHtMltlul, 



Mth-Hihioius l!f. pxfudolmjcUitnfi. anhydmhtosis. microscopy, lauhynides, hirefnuj'ence. 



Introduction 

The tatdigrades or water bears belong to a discrete 
phylum, Tardigrada. ul 1 cosmopolitan distribution 
from diverse habitats including marine. fresh water 
and terreslrud enviioninents. The majority are 
thought lo live m water films surrounding the 
"leaves" of mosses and lichens. I'hey are mieroseopie 
(with adults commonly tanging in length from 200 - 
500 (iltU, are plump and cylindrical in shape a\v.\ haw 
lour pairs ol stubby legs ending in claws. They may 
or may not have eye spots. 

In Australia tatdigrades have been recorded from 
(Queensland, New South Wales. Victoria, Western 
Australia and Tasmania lull not ftom Soillh AliMiaba 
(Melnnes 1994). 

In this paper I report upon the morphology ami 
some aspects of the behaviour ai a tardigrade 
isolated from agricultural sandy loam soil Irom 
Avon, South Australia. 

Materials and Methods 

Loiahts and uiit type 

The tardierades were isolated from a soil of sandy 
loam texture classified as a soloni/ed brown earth 
(Australian suit grouping) 01 as an entisol (US soil 
classification!. The locality was an experimental plot 
on a I arm at A\on (talitude 34' 14' S . longitude 
I3X' 19' it) which was direct drilled and had a 
wheal/wheal rolaiinn. 



I'laslord Road Mitcham S. Ausf. 5062. 



Soil cues (diain. 5 cm. depth 10 cm) were 
collected and mixed in a plastic bag. The sample 
mostly used in these experiments was collected in 
July 1^93 and had been stored dry at room 
temperature for three years. However, frcshly- 
collecled -oil from the same site on 29. tiL 1996 prior 
lo the atiiuiiiu tains was also used lot comparison, 

/.A/mi iioh front .soil 

After thorough mixing of Ihc soil, 50 g altquols 
were placed in a misting apparatus fur ihrce days 
(Ycalcs & Bird 1994). This procedure was replicated 
in t|uadruplicale and, alter three days, the collecting 
tubes were removed and their contents allowed lo 
settle for I b after which the supernatant was 
removed by suction lo within 2.5 em of the bottom of 
the tube. This extraction procedure was used for all 
soils, whether Ireshly-eolleeled of stored. 

Counting 

The contents of each tube were poured, after 
vigorous shaking into a counting chamber I Hone-aster 
1962). The tatdigrades gravitated lo the floor of the 
counting ebambcr between the rings and were counted 
undei a dissecting microscope. 

Ught tninvMop) 

The lardigrades were examined uudei bright field, 
polari/cd light and dillerential inlerlerence contrast 
(Nomarski) optics using a Vano.x Olympus AHBT 
research microscope. 

I iviug tardigr.itles were examined in distilled water 
undcra onershp sealed at its edges with nail varnish. 



I4M 



\ I- BIRD 



Spot- miens were fixed by adding an equal volume 
ol bulling double fclrenglf] h'A 4:1 (20 ml 40',; 
formaldehyde and 2 ml glacial acetic acid in 78 ml 
distilled walei) in a lest inhe to a shaken suspension 
of the i.inti»:rades in distilled wotcr. also in a test 
lube. These specimens were processed lo pure 
glycerol by Seinhorsfs » 1959) method mu] mounted 
hi anhydrous glycerol on slides sealed Lo a eoverslip 
hv molten paraffin as described hy De Maeseneer & 
IVHerde (1963) and then ringed with Lnlclhui 
(Merck). Bolh living and lixed material were 
photographed iw0ng llfonl Ddta 40D film, 

Stftnuini> fleCflQH >ni<Tt>x< <tpy 

lor chsci salions under the scanning cleclron 
microscope fSBM), the fixed material was washed 
lepcutedlv in distilled water, post-fixed and stained 
in l' : < aqueous osmium tetroxidc. washed repeatedly 

again in distilled water, immersed in filtered frcshly- 

made saluraled acjucous Ihioearbohydra/idc lor M) 
min followed by repeated washings in distilled water 
and a repetition o\' the osmium fixation, litis 
osmium-binding technique (Kclley t*t at. 1973) was 
followed hy further washings in distilled water 

Specimens were ftce/c-diied by placing them 
between membrane tillers which were frozen rapidly 
by placing them in a slurry ol freon cooled hy liquid 
nitrogen. The I liters with allached lardigrades wcie 
then transferred rapidly to a freeze drying machine 
and Ircc/c-dricd at -70 ( over a period of Ihree 
doysr. This dried material was Ihen mounted Oil a 
glass covershp allached lo an SFM stub and coated 
with M) nni of gold to enhance stability ami 
Conductivity. I he material was then examined and 
photographed in a Cambridge S 250 Mk 3 SEM 
opcrntcd at 10 k\ using llford 120 roll film iflM 
Plus). 

/, i ,:■!,' i \{), ntnf»f\ 
Altcmpts to determine whether or not M. ef. 



I'sctnlohitji'liunfi would teed on PfltzttetiWto wfanh 
ami thus possibly implicating the tardigrade in 
suppression of this plant pathogen in the field, were 
made using A'. soUfiH grown in Clllttlfc media in IVln 
dishes. Specimens t)l M. Cf, pseiulohnjckmrfi thai had 
been wished repeatedly in sterile distilled watc bv 
centtilugatiou in an elloil to surface slerili/e ihem. 
were poured on lo the fungal plates under aseptic 
conditions in a laminar flow cabinet. 

Results 

Both living and hjtcd I'iciiiieii'-. nt the tardigrade 
isolated from (he Avon soil were examined ^\h\ 
measured. Measurements of the lengths and widths 
ol ten specimens each ol fixed and living material 
showed, as might lie expected, that some shrinkage 
had occurred in ihe fixed material. Measurements of 
the living material were made only on specimens dial 
had their lips and mouth parts retracted sittCC this 
was the state observed in all fixed or dead mateiial. 
The mean length of living specimens was 5 I 1 .4 ± 
47.7 (in) (range 428-580 fim. fable I) and that ol 
fixed material was 423.2 ±48.3 pm (range 364-500 
pm). Similarly, the width oflhe living specimens was 
154,7 ± 15.3 pm flange I2N-I72" uni, Table I) 
compared with }i?1,2± I3.3pm(range IOXI4Xpm), 

The lateral \ tew of the living tardigrade with its 
mouth everted and showing one of each of the four 
pairs of stubby legs ending in claws (fig. I ) together 
with the characteristic inlernal structure of the 
anterior region (big. 2) and the claws at the 
cvlreuulies oflhe fourth pair of legN digs 3, h are 
shown us viewed under the light microscope 

The fixed tardigrade in \cnlro-laleral view with Ihe 
mouth region inverted but showing all lour pairs of 
legs with their claws (Fig, 5i and die ulirastructure of 
the claws on ihe second leg (Fig. (>) are shown as 
viewed under the SFM 



TAlH I 1 \h<<\wri>ici>t\ t>f iivitit; NLiemlmiliis rf pst'iLli-liul. hinli 
Part measured fprm Numbers measured 



I niLili of whole specimen 
VVitllh ol whnlr spei niien 

'■■> petition i>\ stvict 
support on buccal tube 
Length oi buccal nibe 

Wiillh of bueejil lube 

I vngdi ol ph;ii'vn^e;il bulb 

V\ i.lili d| 'pharyngeal bulb 

Length of macroplucoitl 1 1 * 

I I n::tll ul llt.lt roplacoid (2) 
I .cngltl ol mtuupLicoid. 
Length ot 4th (Vint lI.iiw 



Mean 



SD 



Rujilv 



51 14 


±47 7 


428-580 


154 7 


± 15 3 


128-172 


K\M 


± 1 ^ 


7s»..vN2..s 


1(1 ss 


±\.l 


30-42 


5 1 


4<U 


5 5 5 


37.4 


feZI 


35-40 


UA 


±L* 


33-Afi 


10,75 


± i It 


( )-|2 


(.0 


iOS 


5-7 


_"' 


±02 


2.5-3.0 


90S 


±2» 


.-:-■> 



\ TARDIGRADE FROM SOUTH AUSI KALIAN SOIL 



W 



m 



\ 












•■■ 



i S » 



"« - 1 - 



Fig. I . Whole living \hu n>hii>ttts fff, pseudoluifftandi. Bright field optfcS. Lateral view showing everted mil king mouth 

(m), eyespoi te) and one ol each of the lour pain- of legs with lerniinal '.laws (arrows labelled 1-4 from I'roni lit rear). 

The fourth ley is posterior and subleriniiial. Scale ba. = IUO (Jin, 
Fiji, 2- Everted head ol living \f,irmbi<nnx el. pMiiJohuttiuiu/t, Bright field optics showing mouth (ml. buccal tube fb), 

Stytetfi (sj, stylet support (nn). pharynx tp). iTuicrnpliieoids I £ 2 .itnl microplaeoids (arrowheads from top to hotiomi. 

Scale bar = HI (Jin. 
Rig- ^ Posterior rcytoii Of Mucrtthit>m,\ cfi I'sciulolmfvlcmtii. Bright field optics showing claws ol Ihe 4|h leu> (arrows). 

Scale bai = 10 uin, 
Kg -I. Single L'lffW from the 4th leg of Mac)tfhi»nu\ ef t>setuitfltit(il<ifult. Nute hiiMil plate ai base Ofl claw (arrow). Scale 

bar = 10 um. 



LMI 



A. F. BIRO 




A FARPIGRADli -KOM SOUTH AUSTRALIAN SOIL 



LSI 




Pig, 7. Living Mm tohiotus cf. p.wiulohitft-litittti viewed under polari/ed lighl, A. Winkle specimen- Note binTnngence it! 
st) [fibi oo. gill contents and muscle attachments 60 legs (arrow head.-.). Que o! (he iwo eyes fe| is also shown. B Inset 
•Loump an eyp (u) and iln* pronnuneed buelrineener of The Iwo -aylets. Sink bars = SO uni A. 1(1 pin B. 



Although lor general purposes bright liekl opiies 
were (omul to be ntOffi eonvenient (ban fhe oiher 
sv-slonis used, sly lei structure was mueh more 
obvious in living tardigrades under Nomarski :tnd 
poliin/ed light opiies lhan under hrighl field. Stylels 
were, however, clearly observed under all tbree 
oplieul systems provided (hat the htrdigrude was 
alive- In bolh dead and fixed material, the stylels lose 
their integrity and appear lo break down. 

When viewed under polarized light (lag. 7) Ihe 
sislets exhibited marked birefringence indicating a 
regular sirueiural orientation. Ihe slylets are slightly 
curvet), sabiv- shaped si rue lures aboul 40 pin in 
length (Figs 2, 7B). When ihe tardigrades WCK 
subjeeled lo oxygen deprivation on sealed slides, ihe 



stylet birefringence gradually disappeared and was 
entirely lost over a period ol' several hours as the 
stylels broke down. Muscles and intestinal eonieuls 
exhibited birerriugence lo a lesser extent lhan the 
stylels (Pig. 7). However, the muscles and intestinal 
contents retained their integrity and birelnngeuee 
nllcr stylet break down. Six let break down also 
occurred during fixation wliich accounts lor ihn. 
absence in camera lucidu drawings til fixetl material 
m ihe literature. Thus stylets can only be observed m 
living material. 

The buccal tube tl : ig. 2. Table I ) is 40.25 pnt long 
\ 5 1 pm wide (pcrccnl ratio to (he length of ihe 
buccal Uibe(pu - I2.fi7j. The My lol supports th'ig- 2. 
Table It are inserted at JSI'v of the buccal lube 



Lijj. 5 Wlmlc. fixed ^\\i\ •jotd-eonicd specimen ol \ttn'n>hh*\n\ tf. f'.sriu/oiinlrfdiiih, SliM photograph showing tour pairs 

of legs with claws (wtowh. InbHIcil I -J from Ironl lo rear). Scale bai = IHI urn. 
f i<^, 0, SIM photograph ol' a 2nd le*!. Note two *eh ol* double bianehed claws The inner branch in eugjj claw carries two 
it ( r.v. ( >i y |M<tnis (small arrows). Ihe ba,.al plaic ai the base ol the ctaw i-. seen here a*; a rounded thickening (larger 

.nimvJ Ni.ilrlu. 4 |H» 



15: 



A. F. BIRD 



Af*K ■ : 'T* 



<v 



«t 



',-•'-•; 



Y" 7 r 




N 



mc 



Fig. 8 Crushed egg of Macrobiotus ef. pseudohufelandi 
lying within the moulted cuticle (mc). Bright field optics 
showing the "cooling towers-shaped projections on the 
surface of the egg shell (arrows). Scale bar = 10 [am, 



length. The pharyngeal bulb is 37.4 um long x 34.4 
um wide, has apophyses, two pairs of rod-shaped 
macroplacoids and a pair of mieroplacoids. The 
macroplacoids differ in length, the first being 10.75 
pm long (pt = 26.71) and the second 6.0 (am (pt = 
14.91); the mieroplacoids arc 2.9 (jm long (pt = 
7.21). The claws (Figs 3, 4, 5, 6, Table I) have 
smooth basal plates (Dastych & Alberli 1990) and 
well-developed accessory points on their inner 
branches. The length of the fourth foot claw is 8.25 
urn (pt = 20.50). The egg, enveloped by the shed 
cuticle, (Fig. 8) has numerous "cooling tower"- 
shaped projections on its surface which protrude 
about 5 una. 



Taxonomy 

This tardigrade was, using available keys and on 
comparison of measurements, determined as 
Macrobiotus hufelandi Schultze, 1834. This species 
is cosmopolitan (Schuster & Grigarick 1965). 
However, measurements and slides were sent to a 
tardigrade taxonomist (S. Claxton) who kindly 
examined the material and considers that this 
tardigrade most closely resembles Macrobiotus 
pseudo hufelandi Iharos, 1966 and should be 
assigned tentatively to this species (S. Claxton pets. 
comm.). Accordingly, the Avon tardigrade has been 
identified as Macrobiotus cf. pseudohufelandi 
subject to further deliberations by taxonomists. 

Behaviour 

Specimens of M. cf. pseudohufelandi were placed 
in Petri dishes containing the plant pathogenic 
fungus Rhizoctonia solani. Although some of the 
tardigrades lived for several weeks, they were not 
observed to feed, grow or reproduce on the fungus. 
Furthermore, no significant differences were 
observed in the logarithmically transformed numbers 
of this tardigrade isolated from soil that exhibited 
suppression of this fungus compared with the 
tardigrade numbers, logarithmically transformed, 
isolated from soil that did not exhibit suppression of 
the fungus (p < 0.05) (Table 2 ). However, there does 
appear to be a significant inverse linear relationship 
between the numbers of M. cf pseudohufelandi in 
the soil and the numbers of nematodes (p < 0.001). 
Furthermore, a specimen has been obtained alive 
from the soil that was in the process of feeding on a 
nematode (Fig. 9). In soil that had been stored dry for 
three years, there was a marked decline in the 
numbers of nematodes (4 ± 2.9 /50 g soil) compared 
with the number of tardigrades (34 ± 5.8 /50 g soil). 

Discussion 

The first recorded observation of a tardigrade was 
by Goeze (1773) (cited by Nelson & Higgins 1990) 
who referred to them as "little water bears" (Kleiner 



Table 2. The relationships between turdig 
suppressive soils 


rades and nematode* 


/'/; 


Rhizoctonia- 


suppressive 


and Rhizoctonia 


-non- 


So. Is 




Tardigrades 


Nematodes 




Tardigrades 




Nematodes 




Suppressive 

Non- 

suppressive 




Mean 
counts* SD 
19 ±9.1 

32 ±4.8 


Mean 
counts* SD 
401 ± 108 

227 ± 149 




log mean 
counts 1 
2.8! 

3.44 




log mean 
counts' 

5.97 

5.25 




*SED = 65.2 










1 SED 
+ LSD 


= 0.346 
= 0.75 







AlARDIGRAlMil'ROM :i( 



9 




lie y I lead FfcfeO>ii Cri 'i living Mm mhioius at iwudtviuj't - 

Uuuh. Photographed in the process n! fmUny *™ .1 neim- 
tuoV Urigln field optics showing ruuUtfl tm> ami ncrna- 
iiuli- to) Scale hi,. - 111 | mm 



Wvwvt Rijr), Three years lalcr, SpaTUin/ani (J77o) 
iciicd by Nelson & Higgins I Q 00 > called them Slow 
steppers" or // htttli truths which gave rise to the 
phylum intme used today, Because of their 
charactcrislie morphology, the tardigrades (ipc 
recognized today as Kplongine to a discrete p1*> Utni. 
l'hcy probably evolved mine th.in 500 million yews 
ago in the Cumbrian period when there was an 
explosive diversil'ication o(" eukaryoiic organisms. 
Although the Avon tardigrade clearly belong to die 
finji'ItHni) grouping tn the genus M/rro/>/on/v 
iBertoluni & Kebecchi 19951 lis specific la\ononuc 
identity is sub|cel to further deliberation, lliis 
tardigrade hits miiiiI.u measurements to a population 
of /If cl jjstuuiifhttjt^iindi that includes specimens 
from WA (S. Cluxtun pers. conun h Thus, despite Ihe 
lacl Ihal Ihcy have "cooling tower"- shaped egg shell 
projections that are more like Ihose illustrated lor A/, 
lw(?laniii (Nelson & Higgins L99Q) than those 
illustrated lor M. i*stihh>hnjilnnili by Rama//.oin & 
Matkci ( l°N3), which have much broader bases, they 
aiv tentatively assigned to M. ef pStinMHiftifowli (5, 
ClaMort pers. coinm.). 

The ability of tardigrades la survive in a wide 
ranee of environments and their world wide dispersal 
mii-a he due in no small way to their abilitv tp eniei 
inlo an anhydrohiolie slate, a function that they shaie 
with some nematodes and rotifers. I have shown thai 



HIT! I AUSTRALIAN SOU. 153 

M. cl". psaulohujchnult can survive lor at least three 
years in dry soil maintained at room temperature. 
This was the maximum time tested and it seems 
likely that these creatures could survive R>K milch 
longer tinder these conditions, since il has been 
reported (Keilin l$59) that survival times of up to II) 
years can occur. Indeed, recovery- but not survival, 
has been reported to occur alter 120 years of 
anhvdrobiosis (Hancheschi IvMK cited hy Oow-e 
197!}. 

Anbydrobiosis is widespread tn Phylum 
Tardigruda and it is thought that the disucehande 
trehelnse ('unctions to putted these organisms since 
it accumulates within them as they are exposed to 
desiccation (Wesih & RttmAprV IWIJ. In this respect 
tardigrades resemble those nematodes that exhibit 
auhydrobiosis (Madiu &, Crowe (975), hi their 
morphology, of course, lliey are completely ditteieill 
and this is reflected in their different behaviour when 
observed in water under a eoveislip For example, M. 
cf. ptctidrthufeiitndt is able lo push its way through 
an air bubble, This is a capability thai I have never 
observed in a nematode 

The composition of the rwo sabre-shaped stylets 
does not appear to have been studied in detail m<\ 
ihev are absent I'rom camera lueida draw nigs because 
l hey break down and disappear in fixatives. 
Similarly. Ihcy break down on the natural death of 
the animals caused by anoxia on sealed slides. Il 
seems strange that structures which, in the course ol" 
predation and feeding, can penetrate both plant and 
animal tissues, should be so fragile. 

In the living stale, Ihe slyiets are readi!> observed 
undei all optical systems. However, when viewed 
under polar r/ed hglxi they exhibit mailed 
birefringence indicating a regular structural 
orientation. It has been mentioned in the literature 
that the stylets are calcareous (Kaestner l*^S) so 
their birefringence is probably crystalline and then 
break down under anoxic conditions might be due to 
an increased internal acidity leading to the 
dissolution 0[ calcareous structures such as the 
siylets. Clearly such an hypothesis requires further 
testing. 

Mtnri)hii)nfs c[, pscHtlohiijehttuii was not observed 
to feed on the plant pathogenic fungus RfuztnUmht 
Kotimi antl I conclude thai the tardigrade which I 
have follfld in large numbers in agricultural soils in 
Soulh Australia probably does ikm I'eeil on this 
fungus in the Held but preys OH other small sod 
organisms such as nematodes. 

The ability of tardigrades in prey on nematodes has 
been recognized for some Lime. In l°u°- Suyre 
showed that the tardigrade tlypsihins ittyrop.s could 
be cultured using the free living nematode 
himiqrdfit.s rain iru,s as prey fSayre l% l J). 
rurlhermore, Savre {1909) showed that H. myrt'p\ 



1 54 



A I. IIIKI) 



was able lo feed on Ihe plan! parasitic nematodes 
Mtloh/iwytu ftocognUu and uityicnthus Jip.'mci, 
Say re < 1 96V) ) concluded lhal "under certain 
circumstances, (tuitfigntdcs may* give some control 

and these need to be investigated" A major draw 
back to this work was that it was conducted ill a 
moist environment using moss as a substrate. This is 
u lur cry from Ihe normal environment of die plant 
parasitic nematodes tbai Say re used in Ins 
experiments. 

The tardigrade that I have reported upon in this 
paper comes from a line agricultural sod 
environment where it has to survive extremely harsh 
and variable conditions. Under these conditions, it 
may have (he capacity to reduce nematode 
populations although Ihe decline in the numbers ol 
nematodes compared with the number ol tardigrade* 
may reflect the* unhvdrobiotic capabilities of the 
lardi^rades compared with those of the mixed 
nematode population rather than predaiion by the 



tardigi'ades. However, M. cf. pscudoiutft'ltituli \\\.\\ 
be an effective biocontrol agent, although IntJe is 
known of its nematode food preferences and a more 
quantitative assessment of its potential for biocontrol 
of plant parasilie nematodes is needed. 

Acknowledgments 

I wish lo thank Angela Rcid CS1RO Biometrics 
Unit for Ihe statistical analyses. Soils Division 
CSIRO for aeconnnodation, facilifics and expertise 
including that of Stuart McClure for the SEM and 
John Coppi for developing and printing the films. I 
thank Sandra Clayton for advice on tardigrade 
taxonomy and Jean Bud for constructive criticism i)\' 
the manuscript. I am grateful for a granL from die 
Australian Biological Resources Study which 
provided facilities lhal made this research possible. 1 
thank Robin Mauley on whose farm the lardigrades 
wi-iv first discovered lor his kind cooperation. 



References 



lii.iooi \m. K. lV Kihkchi, L. HWt) A revision 0\ Ulci 

Mat -htbt'tHin hnfel<»hli pJUUp (Tardi^iada. Macrobiniidao. 

with some observations on the laxonomie characters tit 

cutaidiiaadcv ZaaL Sttif/ht 22. 127-152. 
Cuowr. J- H ( W1 ) Anhvdruhinsis' an unsolved problem. 

Amur. Nut 105. >b.v 373. 

DASiYCll, H & ArBKiMi. < i (OWO) RedexeripUon <>) 
MttctvhintHs \cmphiius (Daslyeh. J *>7K> comb. nov.. with 
some phylogeuetic notes llardmrada, Macrobioiidae). 
Mitt. humb.'iHfi jWjtf hiM HI, 157-16'). 

Dl MAFSENEER, J. Si D'HliKUE, J. IJV63J Methodcs 
uiilisees pour r etude dps angiulVulcs lihivs dii Rol. Rfvite 

lie i'Asiricutliitr, lirnultex 16,441-447, 
Doncastkk, C, C i 1 962 1 A Limtilina dish lor ncmatodes. 

NnHfttnloyitii 7. 3M-33&, 
Kaisimk. A. ( l%8). "Invertebrate Zoology" Vol, II (John 

Wiley A: Sons Inc . . Now York j. 
Ki n is. [}. (1^5')) The problem of anabiosis or latent lift: 

history and current concepts. Prntu A'r/v. Sot. /.omitm, H 

150, 140 191. 
KltllvK O.. Uikkir, K. A. I & BaiEMIHKj 0. {197JI 

Ligand mediated osmium binding: its application in 

eouklng biological specimens for scanning cfcoirori 

microscopy. ./ l f tlnt,\tnwt. Rrs. 45. 254-25K_ 
M( Inm-s- S- J <l r > l M> /ou»cogruphie distribution of 

terrestrial/freshwater turdij-Tadcs from cunvni literatim.' 

7, Nttt. Hist. 28. 2$7-0ttl 



Mvi.'lN. K- A, ( & CUOWK, ,1. H 1 1975) Auliydrohiosis in 
nematodes, carhnhvdralc and lipid metabolism dunue 
dehydration. .A Exp, /<»<-/. 193. 32I-&HT. 

Ni-i son, D. R. & Hk.oins. K. I'. t IWOlTardiiaada pp, Wi(- 
4IM It, Dindal. DX-. | Ed | "Soil Kioloyy Yn.ide"' (John 
\\ 'dcy X' Sons Inc.. New York). 

Ramwomi. G. Si Mauvi, W, MWO) II Phylum 
lardijiiaila. Tei/a bdi/Joue A/r/ir I\L lini J.iiohtnl. 41, 
1-10)2. 

S.wki-. k, VI, < loon) A method loi euluiring a predutvou> 
tardigrade on the nematode f J unuy,rtlht\ tvtli\'i\>n\. I'uttiw 
Awvn MicWSi'.SSs 21)6-271, 

S< III s|l:K. R. O. & CtHIOARKK. A. A. (1965) r.ildi;4ijda 

from western Worth A merit fl, (hn\\ t'ui t J ith_ AaoL If*. 
1-67, 

Si fNIIORei J W ( 10SO) A rapid method for the transfer ol 
nematodes from tixaiive to anhydrous Ldyeenn 
Nt'nKttoli^tLu 4, 07- 6y_ 

WlMll. V & Ramlov. tl ( l ( WI)Tiehalosu aeenmuhifuHi in 
Ihe tardigrade A,lnnhioiu\ Vtmmff^r durine antivdrobiosi';. 
/ t:\ f >. Zoctk 258. .VM-33 1. 

\\ -vo-s. ('-. W. & Bikii. A. F. f 14441 Seme observation?, 
on Ihe influence ol aunculiuial pruciiees on iln: 
neiualode faunae ol some Soiilh Ausiiahan soils. 
htimhifit ttppi NcimUnl- 17. HI 1 45 



RHOPALOMYIA GOODENIAE, A NEW SPECIES OF 

CECIDOMYIIDAE, (DIPTERA) DAMAGING GOODENIA 

LUNATA (GOODENIACEAE) IN INLAND AUSTRALIA 



By Peter Kolesik* 



Summary 

Kolesik, P. (1996) Rhopalomyia goodeniae, a new species of Cecidomyiidae 
(Diptera) damaging Goodenia lunata (Goodeniaceae) in inland Australia. Trans. R. 
Soc. S. Aust. 120(4), 155-160, 29 November, 1996. 

A new gall midge species, Rhopalomyia goodeniae, is described from stem galls on 
Goodenia lunata (Goodeniaceae) from the Lake Eyre region, South Australia. 
Detailed descriptions of the larva, pupa, male, female and infestation symptoms are 
given. Plants infested by this gall midge are dwarfed and develop few or no flowers. 
Key Words: Cecidomyiidae, Rhopalomyia goodeniae sp. nov., Goodenia lunata, Lake 
Eyre, South Australia. 



tttttlhtottMi "J rht Koutl \oritf\ ,>f'S.Aiht t I9%t Ulli4), 155-lfffl 

KHOPALOMYIA GOODENIAE. A NEW SPECIES OF CECinOMYIIDAE (DIPTERA) 
DAMAGING GOODENIA LVNATA KiOODEMACEAE) IN INLAND AUSTRALIA 

by FfT^R KC)LESIK :: 

Summary 

Koiisik. I! <P'%) Wu'intlotttyut wHklfHUW. a new specie-^ of CeLuloinymlae (Dipteral dam;i£iin> Chnntt'tna 
lunula (Gouuenidceiiet in urUnd Australia. tTrms, A'. Sm . i* , W/. I20(4K |55 U>i>, 29 November 19'Jd 

A new gull midge spcrii-s, tllu>pnhmi\ia & mite nine, is describfd from Sit-in gaUbj on {hnnhniu lunula 
(lioodenueeael from Infl Lake hvre region. Souih Australia- Detailed dextripiiunx of the larva, pupa, male, 
l.-iii.ilr .mhI tnlesi,iiioa symptoms arc glvOW I'laniN infested by ihis j'ull midge arc dwarfed and develop lew Or 
no flower-. 

Krv Woims (Vcnlmiiyudac Rhni>n!om\ia wmJunttu' ffi, IfllYti tj*>t>itvnui hnwut, bake l-~yre. South Australia. 



Introduction 

The insect family Cccidomyhdac is poorly known 
in Australia and Ufltll now. ha* been unrecorded ffaiftl 
the inland regions of fhc coulinenl, The species 
described here was found gulling stems of (Sot nit n to 
Intuitu J. Black iGoodciuaccaei in the vicinity of 
Lake Kyic South Australia in Oelobcr. I99J. 

Soil' <or hairy! goodeuuu Got'ttftflfo htft&tft, '* a* 
perennial herh 5 - 2() cm high with 15 nun long 
yellow Rowers- It grow.s in clay soils along 
walcrcourses and in sandy soils in cenual Australia. 
F'lowering occurs throughout most of the* year but 
peaks between September and November or abcr 
flooding or heavy rainfall (C "unuuiehuiu etf ul ! 1*81 5 
( ooke I9K6). 

The new gull midge species described below is 
placed in Rhoptthnnyin, 3 worldwide genus previously 
known in Australia from only ««ne -species. R. 
taiifornica Fell, introduced lo conlrol Racdutri.s 
Italittufclut (Asieraceac). an American ornameiual 
plain turned pesi on Australian pastures ( Mcl'ayden el 
til, I'W.^Gugnc |9H9a& 

Materials and Methods 

Planls of CiiHHlftttii sp. manifesting stem 
nialFormations caused by Oeeidomyiidae larvae were 
collected al William Creek (50 km west of Lake 
I vo), during a South Australian Museum collecting 
trip, on 22 Oelobcr 19*8; The lew flnweis produced 
by these plants shrank in the course of drying 
precluding luier authoritative identification of the 
species- New plants were sampled into 70% cihunol 
from the same population on 24 August 1995 and 
idcntiTied by D. B. Nyinon, Stale Herbarium of South 
Australia, as G. Itinala. The galls collected on llie 



Department ol I loriieulttire. Vilieulunv and Oenologv 
Umu'iMtv ol Adelaide PMB I Glen Osmond S. Aum. 
SOW, 



firsl occasion were processed in one of two ways. A 
small number was cut open and the larvae preserved 
in HY/< elhanol, A larger number of galls was kept in 
plastic bags and die larvae were reared to adults. 
Pupation look place within the galls Plaslie bags 
were examined daily and emerged adults preserved 
logclher with their pupal skins in ?0ft ethaiiol. 
Canada balsam rnonnis o\' Lypc specimens for 
microscopie examination were prepared according to 
the teehnieiue outlined by Kolesik ( 19451 The type 
series and other material retained in KY't ethanol are 
deposited in Ihc Soulh Australian Museum. Adelaide 
[SAMA| and the Australian National Insect 
Collecriou. Canberra |ANIC|. Dried and presented 
(TO'.i edtanoli samples of mfesled and uninlcsicd 
plants are deposited in the Stale Herbarium oi South 
AuNlralia. Adelaide ISIISA]- 

Gcnus Rhofuihmiy'ui Riibsaamen, 1802 

Rhopalomyia kubsuumen, 1X42: 370 

Type sfwiti's, ()liin>iffn>htts Itttutccttctikt Karsch. 
IK79; Vll.Jber. wcsif. l'ro\ Ver Wiss. Kunsf 27 (des 
Kieffcr. 1896: 89) 

RhcfHiltmiyiu is a worldwide ueiuis comprising 
species ol'Oligolrophiui with t>ne- oi iwo-segmenled 
l^alpi. simple or toothed tarsal claws, elongate but 
entire ci^hdi female abdominal tergite and completely 
seiulose gomtsiylus. 1 he spec ies describe*! here shaits 
with Rhofutlottniii all the above characters wilh die 
exception of luiviiig ihree- or I'our-se^menied palpi- 
As Ihc number of palpal setMnciHs is a derived 
character and varies wilhin several Ohgolroplnni 
genera u does not preclude the new species from 
Rhcpaltiftiyia. 

Rhojwlomyia gootlcniae sp_ nov, 
(FIGS 1-15) 

/A//r'/v/,v rj , William Creek. South Australia |28 



156 



P KOLKSIK 









0$f 



V 



Figs 1-6. Male of Ritnpaiumyia goodenia? sp. nov. I. Genitalia in dorsal view. 2. Sixth flagcllomere. 3. Gonocoxites, 
paramercs and aedeagus in ventral view. 4. Head in frontal view. 5. Wing. 6. Tarsal elaw with em podium. Seale bars = 
100 mn 13, 4; 50 jjm 2. 6; 500 pm 5, 



A NGW M'l < II S-OFCW ll)OMVIII)Al I KUM (i(HH>t.M.\ Lt':%\TA 



1ST 



55' S.. \W 20' fi|, 29.\j,1993, P, fsolcsik, reaicd 
horn larva Irani Ntem gall Dl Qitr/ti&kiU luaala J. 
Black, sampled 22.vl993, r2l32S |SAMA|, 

rV«/y/'<v 2 i3 S, I V/2 pupal skins |SA\1A|. I 6. 
IV, 2 pupal skills |AN(C|, same data; 3 larvae 
|SAMA|, 3 larvae |ANIC|. collected Willi holotype. 

ttth t r material- 2 tfrfc, IV ISAMAJ. s;imc d.»t;K 35 
lu.vac |SAMA|. gal) |SHSA AD995I I27N|. 
eollccled with holotype, 

/Vs. tifttitm 
MnhiVi^ |.f>) 

Colotu: seleroli/ed parts ol body brown, non- 
\deit>U/eil parts of abdomen orange. Wing: total 
lenglh 2.7 mint 2.5 2Xn -4). width 1.0 mm (10- 
I I J; RS at diMal end narrower more weakly 
seleroli/ed mu\ slightly curved posteriorly, joining C 
anteriorly In apeM Rl joining C neat wing mid- 
lengllr Se cell strongly scleroliml and together wilh 
\<\ and adjacent part of R5 bearing scales. 

Hagellomeres 15 in number. Rr,s| and second 
fused, with nodes lunger than necks, thiril to fifteenth 
With nodes and neek> about same length, cireuiutila 
comprising oik IntnsVLr.sL- and one longitudinal 
band. l J alpus lour or thrcc-segnu-nlcd will! well 
developed palpiger in both cases, Tarsal claw simple, 
rounded beyond mid-length, empndin as fanj? as 
claws 

Abdomen; all lergilcs with pair of setae in anterior 
corners, lergiies I VI with single ^cial n>\\ 
posteriorly and a lew seiac laterally- lergilcs VII and 
VIII with 8 few WW scattered m centre: siernitcs II - 
VIII with pair (H s ctae anteriorly, row of setae 
posteriorly and fields of selae both laleiullv and 
central l>. 

(icintaha: gonooo.i|L\s iive venirally. cylindrical. 
Willi small apieoveiilial lube, setose and setulo.sc; 
gonoslvlus situated dorso caudally on goitocosile. 
slightly tapering towards the apex, with apical tooth 
comprising strong claw and a few firm bristles, 
setose dorsal ly ^d setulose throughout: cerci 
bilnbed. wjth several slotti setae on eacb lobe, 
setulose. hypopioet bilobtid, with seta on each lobc. 
,elulose: para meres clasping acdeagtts along their 
full length, setulose. apically bearing lour to hve 
large, setose papillae: aedeagus robust, conical. 

female frags 7-0. I h 

Wing: lola! length | T 9 mm < 1.7 2.1. n i 2i. width 
0,8 mrn (0.7 - 0.9); &5 straight and equally strong 
along lull lenglh as opposed In being nat'iowei aiul 
curved posteriorly ui male Tcrgile VH with selal m>w 
posteriorly, terete VIII with single pair of scute 
anicnorly. sijcrott/aiion ofhodi letvUes in shape ol 
letter "Y". Oviposiloi proirtisiblc, eerei ('used into 



single- terminal lamella, triangular in Uorso-vcuiial 
view, setose and setulose- hypoproct trapezoid in 
dor/so-ventral view, bearing (wo setae posteriorly; 
setuiose. Other characters as in male. 

Ptifhl (Fig. 121 

Colour: non-scleroii/.ed parts ol abdomen orange, 
the rest brown Total length 2.7 mm (2.7 O.8. n = 4r 
Integument of abdominal segments covered with 
spiculae. Cephalic sclerjte with two swelling's as 
long as antennal horns, 30 mn (2S - 36k Cephalic 
pair of papillae Willi strong setae. 148 pm ( 1 4H- 152), 
boms on each side- one of two lower facial papillae 
wilh seta and one of three taicial facial papillae with 
seta. Prothoracie spiracle 92 - 93 pin in length, with 
rraeheu reaching its apex. Abdominal segments I - 
VII with pair of setose vcntial papillae, two pairs of 
setuse pleural papillae, pair or selose and two pairs nt 
aseiosc dorsal papillae. Abdominal segments II -VIM 
doivilly with field of slrong spines on aiHeiior half, 
Abdominal segments VIM and IX with pan ol setose 
ventral papillae, two pairs of seinse pleural pnpill.ic 
and pair of setose dorsal papillae h'aeial papillae 
with setae 5 b urn. papillae on abdominal segments 
wilh setae 5 - H ram. 

Larva (bigs 10. 13, 14) 

Cohan': oran.ue Toial length .3.0 mm (2.6 - 3.4. n = 
fti, Inlcgunieiit covered with dense spicnl.ie, up to 10 
pm long. Head strongly seleroii/ed, 52 ffltl ( 19 5$l 
long and 8S pin iK.s 91) wide, posterolateral 
apodc-mes 75 pm (70 - 79 1 long- antennae 1 7 - 20 pm 
i<>ng and X - |0 pm wade at base. Thoracic and first 
seven abdominal segments w ith pair of ventral 
papillae, two pairs ol'pleurat papillae and llirce pallfi 
of dorsal papillae. Thoiaen. segments with pair of 
sternal papillae and two groups of three lateral 
papillae on each side of -.patula, two of each with 
setae, one without. Abdominal sc-gmt-ni VIII with 
iwn pair< ol ventral papillae, Iwo pairs ol pleural 
papillae and pan of dorsal papillae, all wilh selae. 
Abdominal segment IX beating four pairs ot 
lermuiid papillae, all w:ith setae, bui some papillae 
lacking in sorm- specimens. Amu ventral. Setae on 
slernal and lateral papillae ahoni I um. (»n the olhei 
papillae X - 22 uin, Spatula 294 pin [245 - 32tf) K"ig. 
Wilh apical enlargement I 32 pm (lifi - 1^0) in vvidih 
and 44 pm (34 7 4l<) in length 

hifesfnuan sxmpiotm (1 i^. 15) 

This gj|| midge sjieeies defoims rhi_- stems of 
(toadenia hatata into subglobulai. Iiairv galls, 1-2 
em in diamelei and I 1.5 em in height, flairs are 1-2 
nnii lofl&i dense, grey. The gall consists of many 
globular lo siibglobular. thick-walled cells with one 
larva in each cell. Inlested plants develop lew or no 



158 



P. KOLESIK 




14 \ ® 




® / 



y^ 



A NEW SPECIES OF CECIDOMYIIDAE FROM GOODENIA LUNATA 



159 




Fig. 15. Gal! of Rhopalonnia gooiJcuiae sp. nov. on Goodeiiia hmata J. Black. Scale bar = 20 mm. 



Rowers. On 22 October 1993 at William Creek, 
about 95% of all plants from a population 
comprising some 1000 plants were infested, 

Eivnudogy 
Derived from the generic name of the host plant. 



Discussion 

Currently, the Rhopalomyia genus comprises 86 
Nearctic (Gagne 1989b), 48 Palaearctic (Skuhrava 
1986), 9 Neotropical (Gagne 1994) and I Oriental 
(Gagne 1973) species. No Rhopalomyia have been 
recorded from the Afrotropical Region (Harris 
1980), Until now, only 2 immigrant species of 
Rhopalomyia have been recorded from Australasian 
and Oceanian Regions with R. califomica having 
been introduced to Australia from North America 



and R. chrysemthemi Ahlberg to llie Hawaiian Islands 
and New Zealand from Europe (Gagne 1989a). 
Rhopalomyia is a catchall genus with the bulk of its 
species producing complex galls on Asteraceae. 
Species that form galls on other plant families 
exhibit some morphological differences and their 
placement in Rhopalomyia needs restudying (Gagne 
1989b). The new species described here is the first 
native Australian species to be placed in 
Rhopalomyia and the only gall midge known to 
attack plants of the family Goodeniaceae. The 
species does not breach the current concept of 
Rhopalomyia, except that it has two or three palpal 
segments as opposed to one or two segments in other 
described members of the genus. Although this 
discrepancy precludes identifying the new species as 
Rhopalomyia using the most authoritative current 
key to Cecidomyiidae (Gagne 1981), I find it 
insufficient reason to erect a new genus until more is 
known about its native Australian conveners. 



Figs 7-14 Rlutfuilomvia gootleniae sp. nov.: 7-9. II female. It), 13, 14 larva. 12 pupa. 7. Posterior end of abdomen in 
dorsal view. 8. Posterior end of abdomen in ventral view. 9. Sixth flagellomcre. 10. Head capsule in dorsal view. 1 I. 
Posterior end of ovipositor in lateral view (eerei shriveled in available specimen). 12. Anterior part in ventral view. 13. 
Sternal spatula with adjacent papillae. 14. Two terminal segments in dorsal view. Scale bars = 500 pm 7. 8. 12: 50 urn 
9-11; 100 mil 13, 14. 



Ib<) 



R KOI ISIK 



Rhoptihwyia goodeftfae sp. uov. differs in several 
characters from R. vafifomica, its only Australian 
congener redescribed by Gagne & Boldt (1995). In 
R. goodeniae, the gonosiylus is straight and about die 
same width throughout most of its length, the 
papillae on the male parameres are large ( 1/5 - 1/2 of 
paramerc width), the number of palpal segments is 
three to four and (he antennal horns in the pupa are 
minute and rounded, in contrast. R. califtirrtica has a 
gonosiylus convex at the posterior end; papillae on 
the male parameres are minute (about 1/20 of 
paramerc width), there are one to two palpal 
segments and the antennal horns in the pupa are 
elongate and bifid in frontal view. 



Acknowledgments 

I thank Ifeta and Philip Gee South Australian 
Ouihack Research William Creek for acting as 
guides in die Lake F.yrc region and re-collecting the 
host plain specimens. I am grateful to Mark A. 
Adams and Terence B. Reardon Division of Natural 
Science South Australian Museum Adelaide for 
organizing and leading the collecting trip, David E, 
Symon State Herbarium ol South Australia Adelaide 
courteously identified the host plant species. I thank 
John D. Gray Department of Horticulture, Viticulture 
and Ocnology University of Adelaide and Raymond 
J. Gagne Systematic Entomology Laboratory USD A 
Washington DC USA for commenting on an early 
draft o\' the manuscript. 



References 



( 'OOKE, D. A. 1 19SCO h'amily Goodcniuccae pp. 1383-14 IK //j 
Jessop, J. P. & Toelken. H. R. (Eds) "Flora of South 
Australia. Part 1 (Polemoniaceae - Compositae)" (South 
Australian Government Printing Division. Adelaide). 

Cunningham. Ci. M, Mulham, W. E., Miliiiorpi;, P. I„ & 
I.Hgil U M (1981) "Plants of Western New South 
Wales" (New South Wales Government Printing Office, 
Sydney). 

( i \< .sr. U J. (1981) Ceeidomyiidae pp. 257-292 In McAlpine. 
J, F.. Peterson. H. V., Shewell. G. H.. Teskey, H I 
\mkcioili. J. R. & Wodd, D. M (Eds! "Manual Df Nearetie 
Diptera I" (Canadian Government Publishing Centre. 
Quebec). 

<l9K9a) Family Ceeidomyiidae pp. 152-163 In 
fivenhuis. N. L. (Kd.) "Catalog, of Ihe Diptera of the 
Australasian and Oceanian Regions" (Bishop Museum 
Press and E.J, Brill, Honolulu). 

(l9K9b| "The Plant-Feeding Gall Midge* ol North 

America" (Cornell University Press, lihaca. New Voik). 

( 1994) 'The Gall Midges of the Neotropical Keyioif 



(Cornell University Press. Ilhaea. New York). 

_ & Bonn. P. H. (19953 The gtfll midges (Ptptera 



Ceeidomyiidae) ol tkinluifis spp. (Asterueeaej in the 
United Suites. I>mr. EnUmtoi S»c Wash, *>7<4>. 767-778. 
Harris, K-M.tPJXO) IS, family Ceeidomyiidae pp. 23K-25) 
In Cros^key. R. W. (F.d.l '"Catalogue of the Diptera of the 
Afrotropical Region" (British Museum (Naluml Hislors (, 

London). 

■ 

Kit ii i tfc, I. I. ( IR%> Neue MiUheilungen tlher Gallmucken. 

Wh'fi. Btt.Ztg 13,-85-105. 
Koi.i:siK. P. (I99.S) A new species fff EdchlMicartSa Pelt 

(Diptera: Ceeidomyiidae) on £utj$yplun fasiicuiastt \\\ 

South AuslraJia. J Anst. Int. S/u; 34. 147 152, 
McF.mmN. P.. I.. Dowr 1 1 v. O. P. g TOMliV, A. J. (1983) 

Biological control of groundsel bush pp. 28*30 In Harvey. 

G. J. (Ed.) "Australian Weeds Researeh Newsletter" (The 

Alan Flcteher Researeh Station). 
RiriSA,un n. Ii. I!, ( 1892) Die Gallmucken des Koniglichen 

Museums litr Naiurkunde /u Berlin. fle/A Bit. Z 37. 319- 

41 1, pis Vtl-XVlll. 

SKt iikava. M. (19KG) Family Ceeidomyiidae pp, 72 297 /// 
Sfotii (Kd.) 'Catalogue of Palaeareiie Diptera. Vol. |. 
Seiandae - Anisopodidae" ( Akademiai Kiado. Budapest). 



ROBUSTNEMA FOSTERI SP. NOV., GEN. NOV. (XYALIDAE, 

MONHYSTERIDA, NEMATODA), A COMMON NEMATODE OF 

MANGROVE MUDFLATS IN AUSTRALIA 



By Warwick L. Nicholas* 



Summary 

Nicholas, W. L. (1996) Robustnema fosteri sp. now, gen. now (Xylidae, 

Monhysterida, Nematoda), a common nematode of mangrove mudflats in Australia. 

Trans. R. Soc. S. Aust. 120(4), 161-165, 29 November, 1996. 

A new genus with a single species, Robustnema fosteri, collected from mangrove 

mudflats is described. The circular amphids, six outer labial and six cephalic setae in 

one ring, annulated cuticle, single testis and single ovary with posterior vulva, place 

the genus in the Xyalidae. The buccal cavity is small, conical and unarmoured, the 

lips low 7 and simple. The cardia and ovary are distinctive. Reproductive females are 

wide-bodied with a very large uterus filled with developing eggs and unhatched 

juveniles. 

Key Words: Taxonomy, Robustnema, Xyalidae, Monhysterida, Nematoda, Mangrove. 



KOHVSTNEMA FOSTER! SP. NOV., (JEN. NOV. (XYALIDAE, MONHYSTERTDA, 
NEMATODE), A COMMON NEMATODE OF MANGROVE MUDFLATS IN 

AUSTRALIA 



bv Warwick l . Nicholas 5 



Summary 

Nl( M til At W. I.. ( 199ft) R>>hitstHiiiut [<>\tc'i sp, nov.. guji- llOV: I \v.ihtl:tc Monliystciid:! Nunutud.it, .1 common 
nematode ol inmirrow mudflats in Australia. 7hl/!K H Svi . .V, AlHi/i 120(4). Iht [$?, 2 l ) Nownibci, l'J'J6, 

A new isenvii Willi n single tpectes, Hfflmtftfmu (imerL collected from mangrove mudflats is described The 

Ltivnl.il iimphiiis. six outer labial and ^i\ cephalic MHUB in rmt ring. iinmikiLcil uuiklc, Ringk testis .iuJ single 
iivfli v wnh |hv,u ih>r vulvft, pi. ice the genift in the XyaliJac. The buccal cavily Is mikiII. cortical nruJ imaittnillfviL 
Ihc lipjt low unci simple. The e;inli;i and ovary am tltfittncflVA. Reproductive females aiv wide bodied with a very 
larye uterus Titled with developing cgys and unha<< hed juveniles. 

K i • . woki>\: Tasoiiomv. J\'<>hit\tm-ntu \_wilidac. MouhyMerida. Nunaloda. MangTUYO. 



Introduction 

A new species ol nematode lliat is common 111 
m.iihMint: mudflats in south cast Queensland and 
New South Wales |.s described, Ihc possession i\f 
outer labial and cephalic scum Hue in a single ring, 
circular amphids. an ouisireiched single gonad in 
both sexes, a posterior VlllVU. and an alinulatcd 
cuticle place the new species in the Xyalidae 
| MiHiliysieiid.iK It w'iK previously misidenlified as 
/ Vipjt-vtt sp. tlloclda and Nicholas 1986; Nicholas >' 
ul 1901) because of n resemblance to h. cnuls 
Klome and Schrugc WSJ. However, utotv recoil 
close inspeciion has shown ihal ihc reproductive 
organs are qiuK* clilleicnl and this nematode litis been 
placed in a new genus. 

MiiUrials and Methods 

Type specimens were collected at lOV tide from 
esiuarine mud close in mangroves al Pine River 
F.stuary. which opens into Moreton Hay north of 
Knshane. Additional specimens have been collected 
from mangroves in New South Wales. Nematodes 
were Rxtid in die mud with 5$ formalin immediately 
after eolleelion and were recovered from ihc mud bv 
a combination of sieving and ccntrifugation in 
aq\>COUS colloidal silica (Ludo\. Du Pont de 
Nem(Hirs) with specific gravity adjusted to 1.2. The 
mud was thoroughly dispersed in tap water. 
cenlrifuged at 6000 g for 7 ininules, ihc supernatant 
discarded. Ihc residue re-suspended in colloidal 
silica, eentriluged again at 6000 g for 7 minutes, (he 
supernatanl passed through a nylon sieve with 60 mn 



Division ol BolHHy iiml /uoli^y. Austmlitul N.iU>«nal Ciiner-ny 
(anlvrni ACT 020ft. 



mesh, and nematodes retained by Ihe llher back- 
washed into a pelri dish. Specimens ol the new 
species were picked up with an eyebrow hair 
(mounted on a stick* under a dissceling nucioscope 
and transferred to $% aqueous glycerol. Alter the 
water had been allowed to evaporale al 40 C the 
nematodes were transferred to lresh anhydrous 
glycerol iwm\ mounled on slides with cover slips 
supported by glass beads tballatuui and imped with 
glveeel (CJurr). 

Measurements, given in um. were made from 
drawings of 12 specimens using a camera lueida, 
Type speeimens are in The South Australian 
Museum fSAMA) and their numbers in ihe 
Australian Helminth Collection (AIIC) are given. 

RoSastnemu gen. nov. 

Six inner labial papillae, six outer labial setae and 
six cephalic selue in one ring; circular amphuls. Si\ 
simple low lips. LUiarmoured buccal cavity, cuticle 
aniuilated- Cardia convex cap to anlcrior pair of 
intestinal cells. Single gonad in each sex. vulva close 
in anus, uterus becomes capacious sat holding 
developing eggs and juveniles. 

Rtfbustnemajosieri sp, nov. 
CFIGS I <-.» 

ift>U>t\pc: 6. Pine River Fsluarv Queensland. 
W.vui. l ( )S6. SAM A AHC27695. 
Measurements : Table I . 
f.h'\t riptitm it) v Hi)loi\pt j mule 

Typical nematode form (Fig. 3>, curved ventrally in 
anal region, cuticle uniformly annotated Tail 
proximully conical, posieriad 2Q$ narrow, almost 
cylindrical (Fig. 6». Body setae restricted to five 



I$2 W. 1 . NICHOLAS 

IamiI: I _ MvttsureuwnTs ^/'Rul.uiMiiciiiii foslcri v//, nm: 



Sew 








Males 






Females 




1 V | K 




Holntypc 




Piuaivpcs 


n=5 




Par.it v pes 


lHS 






1352 


Meun 
[242 


SD 


Kange 


Mi-jin 


SD 


Kanec 


1 enjUh 




29,28 


IW-1U73 


I J79 


178 


!0K9-i>v- 


Max. width 




71 


6K 


2. SO 


r,5 7i 


114 


29 


71-161 


C'epluln. flCtnc 


length 


5,1 


5.3 


041 


1.7-5 8 


4 


1.06 


5.5-8.2 


Ainphid ih.nn 




5.y 


7.0 


1 .05 


5.8-8 fi 


5.2 


0.55 


4.5 5,M 


Butt al cavity width 


10 


H,l 


ILS5 


11-12 


12.5 


1 07 


10-14 


Buccal cjvH> t 


upth 


I.I 


IKK 


' 30 


16 :o 


17 s 


2J3 


14-20 


I lead la nerve 


inc 


LOG 


89 


Zl.„1 


54 III 


100 


17 H 


X2-I20 


Phurvu.v length 




212 


224 


MM 


200-254 


: j v-i 


27,0 


102 268 


Head lu vulva 












1098 


170 


S05-i22o 


llcud i(» witty 




1040 


1047 


20,0 


iois nm 


1104 


152 


005-1300 


i.ni length 




22r> 


|*7 


IH.l 


I«5-H2(j 


20f 


17.3 


1X4-220 


Wtdih ai miiuls 




57 


56 


5.27 


(SW>2 


OX 


:• l ! 


50 77 


Spicule, arc lor 


gih 


63 


f>5 


-1,73 


54 Pi2 








ix- Moil's a 




I7_0 


IN 


OOKd 


n.o-io.h 


12.5 


2W 


IMH5.3 


•■ h 




V" 


5.fi 


0.374 


5.0-0.0 


SH 


oj.o 


4 y 6? 


c 




5 5 


6>3 


0.463 


o |.*.7 


7 


0.1.0 


5.0-7.5 


■ c* 




4 


3.5 


i). 120 


Vo-M 


1.1 


4 


■ ft I - 


V'' "■> 












/<) 


A.00 


M ;-, - 



paired, vciiual setae, evenly spaced between anus 
and lail lip, •! pm long, pins three lenoinal 4 pm long 
setae. Six stout lips arched over buccal cavity 
forming shallow dome arising from very shorl 
parallel-sided region, demarcated from cervical 
region by Strong annular groove (Fig. I ). Si\ minute 
inner labial papillae on lips; six short outer labial and 
sin equally short slightly wider cephalic selac 
inserted side by side al base of lips; amphtds circular. 
Situated at level of buccal cavity. Buccal cavity small, 
simple, conical, without teeth, culiculur ridges or 
denticles. Pharynx cylindrical without muscular 
bulb, eardia with about 12 small cells forming 
convex cap to anterior pair of intestinal cells dig. 4). 
No renette cell, excretory' duct or pore, Intestinal 
cells large, paired, with prominent nuclei and 
granular cytoplasm, rectum very short, about M) uin 
long. Testis single, outstretched, to lefl of intestine, 
extending anteriorly almost to eardia. long seminal 
vesicle filled with rounded spcrmalo/oa, extending 
posteriorly U Utile beyond mid-body, long vns 
deferens, very short ejaculaiory duet (Fig. 3). Short 
strong spicules, are length 62 jam (Fig. 7), wilh 
capilulutn, shallow angular bend half-way along. 
exireme lip forms narrow peg: guhernaailum iwo 
slightly curved rods; two very small papdhform pre- 
anal supplements. 46 and 77 um anterior to anus 
(Fig. 6). Three posi-anal caudal glands. 



Paratypes! from Pine River Hstnary Queensland, 
Mviii. loxo, SAMA AiK^/ooo- 27663, 

McLisnremi'nt\: Table I 

I -'i\e ' :* essentially similar to holotypc. The tip ol 
the tail is not necessarily bent dorsally as it is in the 
holotype. Fre-anal supplements vet) difficult to find 
in some males when Ihcy do not protrude in profile. 
Five V V of much stouter build than males, but not 
all arched dorsally as in paralypc illuslraled in Fig. 2. 
Uterus forms a large thin-walled sac occupying more 
than half the body length. The uterus is tilled with 
developing eggs anteriorly and fully -formed 
unhatehed juveniles posteriorly (Fig. 5). There may 
be as many as 28 juveniles and embryonated eggs. 
The egg shell is not rigid and it accommodates the 
shapes of the developing embryos, which are about 
30-40 um long. Juveniles, curled within the thin 
flexible shell are 150-200 \tm long. Three to six 
granular cells, probably unlerlilised oocytes, lie 
along the anterior dorsal wall of the uterus. 

Scanning electron microscopy I Figs K , 9) confirms 
thai there are six cephalic setae, inserted beside the 

corresponding outer labial setae, the cephalic setae 

aiv wider at their bases than the outer labial selac. 
Figures S and 9 show thai the six lips are deeply 
incised and thai the bead is hexagonal when viewed 



Figji.1-7. KflvisttWrnn fasit ii sp. run, I. Mik head 2. Giuvid ten Lite, 3, Fill ire mule. 4. Cuahii. 5, Sni.il I pi mi. mi «4 antiu- 
fcited ankle and uterus containing Unhatehed juveniles 0. Male ijil. spicule*, caudal glands, selac and pir-unal supple 
incuts 7. Spicules and gubcm.ical I. 4, o and 7 holotype male; 2. 3. and 5 paratypc male and len.ialc AH( '276(0 



ROBUSTNEMA FOSTER! SB MOV., GEN. NOV. 



163 




1 n m 5 

JOOfim 6 

lOOum 7 
10 J.im 



64 



W. U NICHOLAS 




Pigs, S-0, Scanning electron micrographs pi head of Rt>hti\tneintt fasten sp. nov. ols outer kibial sola, e« cephalic seta, am 
itinphid, iip fnwr hibitil papiUfl 'p tip. 



ROBUSTNtMA FOSTERl SP NOV, GEN. NOV 



Ifi5 



Differential diagnosis 
The presence of six cephalic setae in the second 
ring of sen si II a, while not unique, is unusual in 
Xyalidae as is the small size of the unarmoured 
buccal cavity and low profile of the lips. The shape 
of the cardia and ovarian development are quite 
distinctive within the Xyalidae. Together these 
characters justify generic status. 



Habitat 
Mangrove mudllats. 



Distribution 

Pine River estuary, opening into Moretoii Bay 
Queensland; Fullerton Cove opening to the Hunter 
River estuary New South Wales and the Clyde River 
estuary New South Wales. 



Discussion 

The female reproductive system is unusual and its 
development warrants further study. Only a 
discontinuous developmental series has been 
observed. In immature females, the outstretched 
ovary is filled with small cells, presumably oogonia. 
After fertilisation, large eggs in early cleavage stages 
appear amongst the small cells. By this time the 
ovary extends forwards almost to the level of the 
cardia. Later the ovary is largely transformed into a 
capacious uterus tilled with developing eggs and, 
towards the posterior, unhatched juveniles. Some 
large granular cells, with single nuclei, lie along the 
dorsal wall of the uterus. These large cells are 
probably unfertilised oocytes and ovarian 
development can be described as hologonic rather 
ihan showing the much more usual telogonie 
development, No spermatozoa were recognised 
within the female gonad. 



Ethnology 

Generic name from the slrongly built body. 
especially the very stout-bodied reproductive female: 
specific name after a colleague. 



Acknowledgments 

I am grateful to The Australian Biological 
Resources Study for travelling expenses. 



References 



Bi.omi: , D. & SaiKAUh, M, (1*985) h'reilebcnde Nemuloden 

aus Antarktis. Veroff. Inst. Mveresforsch, Brenwrh. 212, 

7l-9h. 
HonoA, M. & Nicholas. W. L. (1986) Temporal changes in 

littoral meiofauna from the Hunter River Ksiuary. Attxt. ,/. 

Mar, Freshw, Res- 37. 729-74 L 



Nicholas, W. L„ Elek, A. C, SnSWAftT, A. C. & Maki*ll:s, 
T. G. (1991) The nematode fauna of a temperate 
Australian mangrove mudflal; population densily. 
diversity and distribution, Hydrrtbiotogia 209. 13-2.7 



A REDESCRIPTION OF ASPERSENTIS ZANCHLORHYNCHI 

(JOHNSTON & BEST, 1937) COMB. NOV. 

(HETERACANTHOCEPHALIDAE:ACANTHOCEPHALA) 



By Lesley R. Smales* 



Summary 

Smales, L. R. (1996) A redescription of Aspersentis zanchlorhynchi (Johnston & 
Best, 1937) comb. nov. (Heteracanthocephalidae: Acanthocephala) Trans. R. Soc. S. 
Aust. 120(4), 167-171, 29 November, 1996. 

Aspersentis zanchlorhynchi (Johnston & Best, 1937) comb, nov., occurring in 
Zanchlorhynchus spinifer, is redescribed from specimens collected off-shore from 
Macquarie and Heard Islands between 1986 and 1990. Aspersentis zanchlorhynchi 
can be distinguished from all other species in the genus by having a cylindrical- 
shaped trunk, and proboscis armature of 14-16 rows of 10-12 hooks of which both the 
larger ventral and smaller dorsal hooks have roots. An analysis of the literature 
indicates that the genus Aspersentis comprises four species A. austrinus, A. johni, A. 
minor and A. zanchlorhynchi. 
Key Words: Acanthocephala, Aspersentis, Antarctic, fish hosts. 



JHtntidtWtoU <>f th<> Kovtti .Stn tftx >>l S. Ait\t. ( IW(>). I2tl(4). 167-i !\ 

A REDES1 RIPTION OF ASPERStiNTIS ZAiXCHLORHYWCHI 

(JOHNSTON & BEST, 1937) COMB. NOV. 

(HETEUACANTHOCEPHVLIDAE:ACANTHOCEPHALA) 

by Lhsi.tv R. Smai>s- 



Sumnuiry 




K\ s Wmms; Acanlhneepliala. Ivvawum, Antarctic fish ftc&ls 



Introduction 



Materials and Methods. 



Acanthoecphalan material, collected by the 
Australasian Antarctic L.xpediuon (AAL) ol i u tl- 
1914. was ihc subject of a report by Johnston & Best 
(1037). In rlial report they described hi new species, 
i-tfttHorhyntlms ntmltltfrlnwilii. occurring in the 
stomach of a scorpacnid fish, y,tint'hlf>rhvnt'hav 
\ffiitifer Giiulher. fiWttl Maecjitaue Island. Since (bcir 
description was based on a single female with its 
proboscis not fully everted. Johnston & Rest 1 193 i 
indicalcd lhal Ihc exaimnalion ol additional 
specimens would btf required to eonnnn (he species, 
Ms description and its laxonomic position. 

Subseqnenlly, I wo immature specimens (one male, 
OTW lemalc) weie found m /. spinifi-r collected m 
Maccjuaric Island during Lhe British Australian and 
New Zealand XnUuUie RcfCftKh expedition 
(IMN/.ARFl ol I02°-3I. and identified as I 
:.tinrltlftthxm'hi by Fdmontls ( Im57)_ 

More aeatitlioeeplialans were found when 
members of the Australian National Antarctic 

Research expeditions (ANARE) <>f [#86 - WO 

collected A. .spinifvt (mm Macquarie and Heard 
Islands, humiliation ol this material has allowed a 
more complete description of the atanthoeephnliiii to 
be piepared. ihese specimens, whilst conforming to 
the general desenplions or Johnston & Best ( |037> 
anil l.dmonds (I057) had asymmetrical proboscis 
a i mature and spines mi Ihc trunk, fcaluies 
characteristic ol the Aspersenlinae. The significance 
of Ihese morphological data are considered in litis 
paper antl an analysis of the current status of die 
gCnUS \\pvrscttff\ is given 



'■' I'niversitv "I 'C Vnlial Queensland, Knckhampton Old 

I7te, 



Thirteen /anihlorlivmims spini/i r r collected from 
Macquaiie Muiid waters (M 33 'S. 15* 53'F:)and 
one from the Heard Island shell (trawled between 
51 z 34'and 53 : 30'S. 71 and IK fflTEJ were lived 
m I0 r ; foimaliu Fullered with excess sodium 
tetraborate. Fish were ihen examined under a 
dissecting microscope and any aeanihocephalans 
found were stored in 70'.f elhauol prior lo 
examination, either as temporary wet mounts, after 
clearing in becchwood creosote, or as permanenl 
preparations, after staining in Grenadier's carmine 
alum, dehydrating through a graded series ot elhanol. 
clearing in xylene and momiLing in Canada balsam. 

Measurements o\ 10 males anil 10 females were 
made with the aid ol an ocular micrometer, drawing 
tube and measuring wheel and are given in (.tin unless 
otherwise slated, with the range followed by the 
mean in parentheses. Figures were drawn with (he 
aid o\' a drawing lube. All specimens have been 
deposited iq the Queensland Museum (QMi 

Systematic* 

Order Palaeaeanthoeeplnila Meyer 1031 

family lleteracanthoeephalidae PeirochenU), M?5fc 

Subfamily Aspersenlinae Golvau. IvbT) 

Genus A\(H'r\eitti\ van Cleave. 102° 

Type species AsfH'rtcnlis tttittHnu\ Vail Cleave. 

Kspersentis zanchlorhymhi (Johnston <& Rest. 
1437* comb, nov 
(FIGS 1-7) 
Synonym EilitntHitvucltax :,Liin'hh>rh\nthi 
Johnston & Best, 1037 pp. 12 L& Fdmunds, 1057 p 
96, /-,v/, .-;f/^c///^r/M^rA//d/itowies.|M, 108S p{>. NO. 
I0J. tabic t. 



IbX 



I.R SMALBS 



Of^- 




Figs 1-7 Asf>ersen(i\ T.aiiclilorhynciii comb, nov- Fig. I. Proboscis armature, one row ot'dorsnl hooks. Fig. 2. Proboscis 
armature, one row of ventral books. Fig. 3j Proboscis, dorsal view. Fig. 4. Male. Fig. 5. Female, proboscis not fully evert- 
ed, showing distribution of trunk armature. Fig. 6. Posterior end, female. Fig. 7. F^gg. Scale bars = SO uni. 1. 2: ISO urn. 
3i 100 um.Vv 500 um. 4. 5; 25 urn. 7. 



A REDESCRIPTB IN < >f tAH -ftsi -\>us / \\>( /// <M7/> ,v< w 

MtHcriu/ examined 
From /itih-lt}i>r!mt<hn\ spiiufer. 126 ? 9. 170 



'," 



n'rj. Macc|uaric Muiid, 6.H.86. l2.vi.S6, (vxii.SO; 
02 J 1324^3211335. IV Heart Wand, ILviOO; 

(;:i 1333 

Revised de\eripti'm 

Trunk cylindrical Pmboscis long, cylindrical, sel 

at angle to trunk i^te* **» 5) Protects atmuunc 

similar in both sexes, 14-16 fOWp Ci£ 10-12 hooks 
(Fig. 3); dorsal rows of hooks (Kit;. I) somewhat 
smaller than ventral rows (Fiji 2): all hooks with 
roots. Neck shorl unarmed, ttuoeulcd, Spines tiny, 
einhedded in cuticle in both sexes, encircling anlerior 
emi iri Hunk to A tcV6l abOW halfway down proboscis 
receptacle, then extending down lateral Irunk 10 
postetior [Fig. 5)1 Proboscis teceplaele double- 
walled, inserted at base of proboscis; ganglion 
placed near posterioi eml, Lcmnisci flat, longer than 
proboscis reccplacle when fully extended, 

Male: T \i 3 T 4>6 <4.2) mm long by 360-6K0 

(180) wide. Proboscis, nol fully extended in plOSl 
specimens. 650 long by 215 wide (n-lf Largest 
dorsal hooks M\\ and -lib in row, S3 fi& largi_>i 
ventral hooks Vd and 4lh III row 76-85, Neck I3CT- 
|95 (145) li>ni> by 140 260 (175) wide. Proboscis 
receptacle 615-99? (735 1 ling, lcmnisci 741) 1 300 
(930) long. Testes ovoid. Utndeinly placed; anleitoi 
testis 455-9S5 (075) long bv ISO - 300 (265) tvitfc 
posienor testis 490-7 1 5 (635 ) long by 195445 [280] 
wide. Cement glands, six, pear-shaped. Male 
aperture terminal. 

Female: Trunk 5.3-16 ( I0)mm king by 300-765 
(500) wide Proboscis^ not fully extended, longer 
than ''00, width 200. Largest dorsal hooks. 3id and 
4th in row. 50-56; largest ventral hooks. 3rtl and lib 



in row 80-83. Neck 182-227 (200) long by 175 200 

(205) wide. Proboscis reccplacle SI0-I300 MIOOl 

long, lcmnisci 925-1940 long (n-2), Female aperture 

terminal (Fig. 6). F.ggs embryonaied. with 

prolongations of middle shell 75-90 (87) long by IS 

wide (Fig. 7). 

Ifost: '/nth hltirhxnclttis spittifer Giinllici. 

Location; stomach, intestine. 

Locality: Maeciuaric Island, 

Type specimens: Hololvpc female. South Australian 

MuviiinV J200, 

Remarks 



Alihough a large number of specimens was 
collected, none o\' them hud been relaxed and 
extended prior to fixation, which made them difficult 
ti> study. Comparison of the material from this study 
With the dcscriplions ol Johnston & Best < PU7) il "d 
Edmonds 11*357") showed lhat all the material 
collected from /. \pinifcr was the same species. In 
many specimens the lcmnisci were as described by 
Johnston & Best ( |93?J that is, shun and irregular 
and reaching aboul one-third ihc length ol the 
pmhosiis receptacle but in I he more relaxed 
specimens, the lenuiisei were flat and extended 
beyond the proboscis receptacle. The cylindrical 
shape of the proboscis and it* armature, 14 In rows 
ol 10-12 hooks, were Observed in the three 
specimens described by Johnston & Best ( 1 937) and 
l.dmonds (1957) but in none ol them had the 
proboscis extended far enough to describe the 
morphological details Of the proboscis hooks. The 
asymmetry o( the armaUuc. ventral hooks being 
larger than dorsal ones, could be >een onlv in those 



lAIII.I. I. /\ .ctttHftttfiMW (if foliate h t ulv n><\t,',ttr<>m<'iit.\ <>/ Aspeisenli* auMiiiius Vim ( leave, 1^29 (taken j ami 
/,i;i!<ns-ic,-U I'JSh.A. minor fithtwnJs S: Swales. IW? aint A. /anclilnrliMichi iJofmstan i< Best, W7). Measurement' 



A, nustfvtus 
South Sheiliinti-, South Georgia 



o_94-X.54(7.25) 
(J»-2.UU(|.73) 
0.07-0.73 (0.70) 
0.294 >_35 1(1.32) 



trunk length 4.93-6,42(5.79) 

trunk width LI6-1 .79 ( i .19) 

prohmeis lei^lli 0.51 0.66 (0,5O> 

proboscis width o 294U2 0L3O) 

dorsal hook length f maximum! 0.054 0.0h4 (0 060) o 00O-O6S0 (0,063) 

\L-iunil iMiok length (maximum) U.I 194). M/ (0.126) 0.132-0, I4 1 M0. 1401 

neek length If 17 23 (0.I26» 0.22-0.31 (0.27) 

egg 0.0604U1NX 0.07141.0X7 

\O,0|U4>,025 H 02041.025 

lio4ikdis|KtMlion IJld BOW* of 7-1 1 llooks/ro\v 



■\. rniH/n A. zoin (ilorhytichi 

Tusmania Maequarie Sc Heard Is 



Z3 4 i i 3?i 
(i.M MS Ut.s4i 
0.24-0, ^i().2S) 
0. 10-0.17 (0.14) 
O.03(M).035 i(L032) 
0.06M),OKO (0.065) 
0.12-0.25 
0,06M4).077 
\0,IH2-0_()lfi 
14 rows of 7-9 
hooks/row 



5.VK..0ll0.0» 

0.39-0.76 ((150) 

>0^0 

0.20 

0.050-0.050 
0.OS0-0.0S3 
0.164.23(0.201 

0.07s (),0<HM0,08?) 

k (LOIS 

14-16 rows nl 10-12 

houWit^ 



171) 



LR, Smai.i.s 



specimens with the proboscis almost, or completely, 
everted. This character is indicative of the genus 
Aspersentis rather than the genus Echinorhynchus to 
which the species was originally allotted. Somatic- 
armature, present in this species, is also found on 
other species of Aspersentis (sec Zdzitowiecki 198 1. 
1986) but not Echinorhynchus. Since liny spines arc 
easily overlooked, as has occurred in some 
col lections of A. austrinus (see Zdzitowiccki and 
Rokosz 1986). it is not surprising that they were 
undetected in the earlier studies. 

Aspersentis z&nchlarhyncfii (Johnston & Best, 
1937) comb, nov, can be distinguished from ,4. 
austrinus Van Cleave, 1929 in having a more 
cylindrical trunk, a longer proboscis, at least 650 in 
males and 900 in females, compared with up to 630 
in males and 720 in females, in the distribution of 
trunk spination on the lateral trunk as well as 
encircling the anterior trunk, and less marked 
asymmetry of the proboscis armature with both 
ventral and dorsal hooks having roots. Aspersentis 
zanchlorhynchi has more hooks per row (10-12) than 
does A. austrinus (7- 1 1 ) on the proboscis (Table 1 ). 

In comparison with A. minor Edmonds & Smales, 
1992. females 3.2 mm. and A. johni (Baylis. 1929) 
Chandler, 1934, females 3.0 mm, A. -.anchlorhynchi 
is much larger, females 10 mm. With 10-12 hooks 
per row on the proboscis. A. zanchlorhynchi has 
more hooks than A minor, 7-9, and fewer hooks than 
y\, johni, 1 2- 1 4. Aspersentis minor occurs in 
Rhomhosolea tapirina from Tasmanian waters 
(Edmonds & Smales 1992) and A. johni occurs in 
Merluccius sp. around the Falkland Islands 
(Yamaguti 1963). By contrast A. zanchlorhynchi 
occurs in Z. spinifer from Macc|uarie and Heard 
Islands, that is from sub-Antarctic and Antarctic 
waters. 



Discussion 

In a redescription of A. austrinus Zdzitowiccki 
(1981) placed Rhadinorhynchus wheeled Baylis. 
1 929, Aspersentis wheeieri Chandler, 1 934 and 
Aspersentis megarhynchus (Linstow, 1892) Golvan, 
1960 nee Echinorhynehus megarhynchus Linstow, 
1892 as synonyms of A. aitstrinns. He commented 
that Linstow (1892) gave the number of proboscis 
hook rows as 18 and described the trunk as unarmed 
and that Linstow neither measured, described nor 
drew the ventral and dorsal rows of hooks as having 
different shapes and dimensions. All of these 
characters arc inconsistent with the genus 
Aspersentis. Amin ( 1 985), however, in his 
classification of the Acanthocephala overlooked 
ZdzitowieckLs paper and followed Golvan ( 1960) in 



listing A, megarhynchiis (Linstow, 1892) with A. 
austrinus as its synonym and A. johni (Baylis 1929) 
as the only two valid species in the genus. 

Zdzitowiccki & Rokosz (1986) rc-evalualed die 
validity of Heteracanthoeephcdus hureaui Dollfus. 
1965 and concluded that it was either a synonym of 
A. austrinus or. because of the wide range of number 
of hooks per row. of A. johni. Zdzitowiecki ( 1 986) in 
his systematic review of Antarctic acanthocephalans 
reaffirmed his conclusion that Echinorhynchus 
megarhynchiis Linstow, 1892 did not belong in the 
genus Aspersentis, and listed H. hureaui as a 
synonym of A. austrinus. Then Zdzitowiccki ( 1990), 
when re-examining material previously designated 
//. hureaui, staled that, "as was suggested earlier H, 
hureaui is identical with A. austrinus = A. 
megarhynchiis". Edmonds & Smales (1992) noted 
the inconsistencies in the designation of A. austrinus 
by Zdzitowiecki (1981, 1986, 1987, 1990) and 
indicated that E. megarhynchiis as described by 
Linstow ( 1892) lacked body spines and did not have 
asymmetric proboscis hooks. 

Thus. A. megarhynchiis as redeseribed by Golvan 
(I960), is not the same species as E. megarhynchiis 
Linstow, 1892 because it has asymmetric proboscis 
armature and cuticular spines on the trunk, and is 
now considered as A. austrinus. Since A. austrinti\ 
was described in 1929 by Van Cleave while A. 
megarhynchus was not established until 1960 by 
Golvan, A. austrinus becomes the type species of the 
genus Asftersentis with ,4. megarhynchiis as a 
synonym. The other synonyms are A. wheeieri 
(Baylis, 1929), Rhadinorhynchus wheeieri (Baylis. 
1929) and Heteracanthoeephaius hureaui (Dollfus, 
1965). 

The genus Aspersentis therefore now consists of 
four valid species A. austrinus, A. johni, A. minor and 
A. zanchlorhynchi. 

As to Echinorhynchus megarhynchus, Johnston & 

Best (1937) suggested that it might be identical to 
Leptorhynchoides debenhami (Liepcr & Atkinson, 

1914) Johnston & Best, 1 937 now Metacantlux rphalus 
rennecki (Liepcr & Atkinson. 1914) Zd/ilowiecki. 

1 983. Echinorhynchus megarhynchus occurs in Notothenia 
corriceps, one of the hosts of Metacanthocephalus 
johnstoni Zdzitowiecki, 1983, but not in Trematomus 
hemachii the host of M. rennecki (sec Zdzitowiecki 

1983). Moreover the original description of E. 
megarhynchus by Linstow (1892) resembles that of 
M. johnstoni in having a proboscis armature of 18 
rows each of 6 hooks, the proboscis 0.45 mm long, 
and apparently no neck. Of the other species 
occurring in Antarctic fish hosts, Echinorhynchus 
petroschenkoi (Rodjuk. 1984) Zdzitowiecki. 1989, is 
a larger helminth than E. megarhynchiis with a 
longer proboscis, proboscis armature of 15-19 rows 



A RUDHSCRIPTION OF ASPERSENTIS 7ANCHL0RHYNCHI 



i ;i 



Qf 10-13 hook*, ami docs not occur in N. corhicvjKs 
(see ZdZitowiecki 19K9K HeterosenTts heteracanthus 
(l.inslow. 1896) has body spines and proboscis 
armature of only 10 rows of 4-5 hooks, with a 
striking difference between the length of the first two 
and the last three hooks (Zdzitoweicki 1984). These 
characters suggesl that E. ttiegarhvnchtis is closer to 
M. johnsumi than any of the other aeanthocephalan 
species occurring in Antarctic fish. Direct 



examination of specimens of A-/, johnstmti is needed 
before a determination on the status of E. 
nH'garhynvluis can be made. 

Acknowledgments 

My thanks to Dr R Williams who collected the 
parasites and Professor K. Kohde who made them 
available. 



References 



AMJN, 0, M. ( 1085) Classification pp. 27-72 hi Cromplon. 

I). W. T. & Niekol, B. B. (Eds) "Biolopy of the 

Acanlliocephalu" (Cambridge University Press. 

Cambridge). 
EliMQNDS, S. J. (1957) Aeanthocephala. B.A.N.Z.A.R.E:. 

1929 -31. Report ser B. 6, 91-98. 

& Smai.i-:s. L. R. (J9921 A new species of 

Aeanthocephala from the Greenhaek Flounder, 

Rhomhasolea tapirina Giinthcr. 1S62. Trans, R., Soe. S. 

Aust. 116,35-38'. 
GOLVAtt, Y. .1. ( 1 9nO» Le phylum Acanlhocephala. 

Troisicmt* note, La classe des Palaeacaulhoeephala 

tMcycr. IWh. Annfs. Parasilat, hum, camp. 35. H8- 

1 63. 
Johnston, T. II. & Rusi; K W. (1937) Acanthocephala. 

A.A.E. 1911-14. Report ser. C. 10. 5-20. 
Lin.stow, 0. von 1 1X921 HelmJnlheii von Siidgeorgian. 

Nach tier Ausheute tier deulsehen Station von 18R2- 

l*K3../u/ir/>. flamh. wissett. Aust. 9. 10-77. 
YAMAoini. S. (1963) "System;) Helminlhum" Vol. 2 

Aeantlioccphala (John Wiley & Sons., New York.}. 
Ziv.itowii.ckl K. (I9K1) Redeseription of Aspersetuis 

anstrinns Van Cleave, 1929 i Aeanthocephala). Acta 

Rarasttol, Pol, 28, 73-83. 



(1983) Antarctic aeanthoeeplialans of the 

eenus MefacauthocepJuilns. /hid. 2X, -113-437. 

( 19S'4) Dcscnplion of Heierasentis 

hew meant has (Linstow. ] S96) from Antarctic fishes, anil 
remarks on the laxonomie status of Hftewsemia Van 
Cleave. 1931 ( Acanthocephala. Arylhmacanlhidae). Ibid, 
29. 11-115. 

1I9K6) Acanihoecphala of the Antarctic. 

Polish PoL Res. 7.97-117. 

(1987) Aeanthocephalan* of marine fishes in 

Ihe regions of South Georgia ajid South Orkneys 
(Antarctic). Aeta Parasitof. Pol 30, 211-217. 

( 1989) A rede.seription of Echittarhymhus 



petrotschenkoi (Kiuljuk. I9&4) comh. u. (Aeanthocephala). 
Ihiii 34, 1 73- 1 SO. 

(1990) Rc-exauiinaiioii of five Antarctic and 

suhantaretic diyenean and aeanthocephalan species from 
Professor S/idaCs collection. Witt, 35. 31 36. 

& Rokosz, B. ( 1986) Prevalence of 

aeanthocephalan s in fishes of South Shel lands 
(Antarctic). II Aspersenfis itusthnns Van Cleave. 1929 
and remarks on the vulidil\ of Heieimanthoecphalus 
hureaui DoWWis, 1965. 1 hid. 3$, 161-171. 



SEASONAL ACTIVITY OF THE EARTHWORM, 

GEMASCOLEX LATERALIS (MEGASCOLECIDAE), IN A 

EUCALYPTUS WOODLAND IN SOUTH AUSTRALIA 



Brief Communication 



Summary 

The earthworm fauna of agricultural habitats, especially pastures, has been 
extensively surveyed in south-eastern Australia in recent years 14 . The fauna is 
dominated by accidentally introduced species, particularly Lumbricidae from Europe 
(e.g., Aporrectodea caliginosa, A. trapezoides and A. rosea). Native species are rare. 
Very little is known of the biologies of native earthworms, either in urban, 
agricultural or native habitats 3,5 n . This brief communication reports on the seasonal 
activity of Gemascolex lateralis (Spencer 1892) (Megascolecidae), one of the most 
common native species in South Australia, and offers one possible explanation for its 
rarity in pastures. 



Tmnsactiaw of the Srryal Saviely W s' fowl- 1 Wok 120/4). 1 73-174. 
BRUT COMMUNICATION 

SEASONAL ACTIVITY OF THE EARTHWORM, GEMASCOLEX LATERALIS 
(MEGASCOLECIDAE), IN A EUCALYPTUS WOODLAND IN SOUTH AUSTRALIA 



The earthworm fauna of agricultural habitats, especially 
pastures, has been extensively surveyed in Mouth-eastern 
Australia In recent years 1 " 1 . The fauna is dominated by 
accidentally introduced species, particularly Lumbricidae 
from Rliropc (e.g., AportVCtodeii caiii;ifu>su, A, (iu})c:tiith's 
and A. rplnsd), Native species are rare. Vei> little is known 
of trie biologies ol native earthworms, either in urban, 
agricultural or native habitats' ? ". This brief 
communication reports on the seasonal activity of 
(,tMit\<<>U>\ laicralis (Spencer 1892} (Megascolecidae). 
one of the most common native species in South Australia, 
and offers one possible explanation for its rarity in pastures. 

Eneclbrook National Trust Reserve is an open forest 
iEutrih'pitts uhlicfmi - E. hoxteti) at Hridgewater in the 
Mount LoJiv Ranges, SA (350 m altitude. Grid reference 
962224 on' Sheet 6627-1. Echunua, I ; SOQOd), The 



understorey of the reserve is composed of shrubs such as 
Banksia, Acacia, HuU-a, LcptospertmuH and Hthhvn'ta spp. 
The soil is a yellow pod/olic Average annual raintall is 
1050 mm. Much of hngelbrook Reserve has been burnt hy 
busiitlres in recent years but the sites reporled here had not 
been burnt for at least 35 years at the lime of the studv 

Baker' : ' ' set eighty pitfall traps (plastic jats. l > cm diarn,, 
9 cm deep) flush with the .soil surface within l:ngelbrook 
Reserve in March 1983. Forty (raps were set on a north- 
west slope and the other forty on a south-west slope. The 
traps were set 10 in apart in two transeeLs on each slope. 
Fach nap was covered by a ceramic tile (15 s 15 cm), set 
appioximaiely 2 cm ahove the traps on three nails. These 
tiles prevented rain and leafliuer from fouling the traps. No 
preservative was added to the traps. The traps were 
inspected weekly until March 1MN4 and the invertebrates 




A M J 



J 



SON 



1983 



I 

i 


1 


si i 

mmJL 


i \j ii 1 1 1 1 1 1 1 1 1 

D J F M 






1984 



120 



100 



80 



20 



m r\ 



E 
E, 



60 -s 



40 



(0 

»— 
c 

03 

CD 

CD 



l»g. I . Numbers CtfGtvWCaleX laWralh trapped per week in Rngelbrook Reserve, S A during I983-K4 (bars). Weekly 
rainfalls, recorded nearbv at Stirling Post Office, are included (line). 



174 



UK IMKF.R 



thai were cmieht Were preserved fur later counting. 

One species of earthworm. (V. lawnilis, was trapped, 
mostly in the summer months (Fiir 1). There was n« 
discernible difference helwecn the catches on the ivvu 
slopes H = 1.42. p > 01)5 lor a paired t test comparing total 
weekly cinches tor the two slopes), the data lor the two 
groups of traps have therefore been combined in \ s \\*. I. 
There was no apparent relationship between the peak in 
napped eaithwoims in December and prevailing weather 
(c e.. tahifal)) at thai lime (r = - 0.239. /> > 0.05 lot wceU> 
data tor 9 NWmlvr l*>H3 to 21 March 1084), 

Tlie:iLti\i lies ul earthworms, on and near the soil surhu.e. 
are usually associated with coo], wet weather in temperate 
and mediterranean habitats" * In particular. Abbott 
reported that the activity of native earthworms in a jarrah 
lotest in south-western WA was restricted to wmtei and 
spring when -oil moist uic was highest, It is therefore 
sin posing thai the surface activity df G. ItitvmltA peaked m 
stunmcr at Kngelnrook Keserve when weather was at its 
hottest and diiesL However, Lawson 1 " found (i. hucrnti; 
Ihrouphout (lie year in the surface Livers ol the sod (0o mi 
Jeep) In U kutvlyfKUi woodland near Cape .Icrvis, SA. She 
arjiued llial. hecause ol the predominance ol elitellate .utoh* 



at this time, (V. Inhnifis reproduced during the hotter, 
wanner months o| (he year in this woodland. She did not 
find cocoons (at any time) lo vrnl'v this conclusion. In 
addition. l.awson could not find Q. lateralis ill a nearby 
pasture. Baker eM/A" did find siiull uinnhers of 6'. htlvmlis 
in other pastures in the Mount Lolly Ranges, In ihese 
pastures. G. laUralts was present in the top 10 liii ol soil 
from autumn to spring but retreated to grcak'i depths in the 
sammei mouths. Most worms were found in a patch within 
one pasltirc near a clump of Eucalyptus tices .md lallen 
log*, 

Why C talemtiji is present, indeed most active, on the 
soil surface in native woodland $ during summer is not at all 
clear. Apparently, the shelter provided by die above-ground 
vegetation in woodlands provides sufficient nioi-.tnie jnd 
cool temperatures lo enable such behaviour hint in open 
pastilles, it seems likely that the lack of similar vegetation 
would prevenl this summer surface activity and heme 
reduce the abundance ot (', Utirnili\_ II i| js to survive in 
pastures at all ihrotigh summer. G htioralis musi retreat 
bom surface layers to greater depths when- conditions jje 
Lonlpr and moisler. as do the more abundant, introduced 
lunibiieids IJ 'v 



Baker. G.H., Buekerlleld, J.C., GrevCuidncr, l<„ 
Merry, R. & Doube, B.M. ( L992) Soil Biol, Biochem. 24 

1 38*>-I395. 

Baker, C.I!., Barrett. VJ., Williams, P.M.L.. Curler, Ej. 
& liuekerlUld, J.C.I )*)93) "Distribution and abundance ol 
rMiihworms in south-eastern Australia ;mm\ their inlhieni e 
on the burial ol lime" pp 145-348 /// Corey. S.A.. Dull, D..I. 
& Milne, W.M. (Ed* J "Pest Control and Sustainable 
Agriculture" (CSIKO. Melbourne). 

Baker, G.H-, Thumlerl, T.A., Meisek L.S., Carter, P.J. 
& Kilpin, G.P. ( 1996) Soil Biol, Bioehem. In Pre** 
'Baker, G.H. (1996) "The ecology, man:igemeut and 
benetiis nf earthworms in agricultural sods, with particular 
nTenmcc to southern Australia In Bdwards. C.A (Hil.t 
"Earthworm Ecology" (Soil and Water Conservation 
Society ol AH1C11C& Ankcuy). In Press. 

Wood, ISO. ( 1974) J. Ariim, l.col. 43, N7 1 0b. 

Ahboll. I. t 1985) Aust, .1. Soil Kes. 23. 263-270. 

Abboll. I. ( 1 985 ) Ihiil. 23, 27 1 -290. 



Tan Praagh, B. (1992) Soil Biol. Rtocltem. 24. MM 
1367. 

Baker, G.H., Barrett. V.J., Grey -Gardner, R. & 
Bucket-Held, J.C. (19931 Trans. R. Soe. S. Aust. 117. 47 

' Lawson, L.M. (199& The Disiribuiion and Abundance 
ol Native and Introduced liarthvvotms in an Area ol* Pasture 
and Native Vegetation near Cape Jervi.s. South Australia" 
PhD Thesis, Hindcis University of South Australia. 

'Abbott* 1. 1 1994) Aim. I, Sod Kes. 32. I 17-126. 

Bakei; G.H. ( I9S6) lians. K. Sot. S. Aust. 1111, 4MS. 
1 'Baker. C.H. (l^KK) Aust. l-nl. Mae,. 15. l27-t39. 
"Kee, K.K. (l c J85l "Earlliwtirm^. Their tieoloey ajill 
Relationships with Soils a\u\ Land Use" (Academic Press, 
Sydney). 

Tlouehe, M.B. ( L972) "Lombriciens de France, bcolo^te 
et Systcituilique" (INKA. Paris). 

Baker. C.H„ Crey-Cardner, R. & Buekerllcld, J.C, 
(IW)Ausl. J. Lcol, 17, 177-188. 



( .11 BAKER, Division ot Entomology, CSIRO PMB 2 Glctl Osmond S.Ausl. 5064 



GASTROINTESTINAL PARASITES OF FERAL CATS 
IN THE NORTHERN TERRITORY 



Brief Communication 



Summary 

The feral cat, Felis catus, is well established as a predator in Australia and feeds on 
insects, fish, amphibians, birds, reptiles and native and introduced mammals 1 . Several 
surveys of the gastro-intestinal parasites of feral cats have been carried out in south- 
eastern Australia 2,3,4,5 , the species and prevalence of the parasites encountered varying 
between states, depending on available food sources and climate. In this study we 
present information on the gastro-intestinal parasites recovered from feral cats 
collected from the Northern Territory, a region from which only limited data currently 
exist. 



Tninwtuw t*f the Ht>\vl .W/.*/\ ttj tS Arm llttft) 1 20(4 1. 175 I 70. 
BRIEF ■" COMMUNICATION 



G AS I 'RO-INTESTINAL PARASITES OF FERAL CATS 
IN THE NORTHERN TERRITORY. 



I hfl feral till. /('/'> I atti\. is well CStqbHHltcJ as N pK-ditUti 
in Australia mo! lefils on iiuects, [jib. amphibians. fcfirUs, 
reptiles nil J nulive am' introduced mammals' SiSVfifftl 
surveys ftl ilu- gasiro-intcslinal parasites of I'cral Cflh hwa 
been carried mil in south-eastern Australia 1 ' ''. the .species 
and prevalence o( Ihe parasites encountered . ;.r\ «««g 
between states. dependim 1 . on available lood mhiivc* ami 
c1imulc_ In Lifts xivitly WC present information 013 tbe unMn«- 
intestinal parasites recovered liom lend cats collected ftftffl 
ihe Northern lerriiory. a region IWittl which only limited 
ilaia turivuilv cxifti 

Stall liom Ihe Parks ;mkI Wildlife CflMDltMfijftO ill 'he 
Northern Teoitory napped and/or shot I'ftfi feral eals foi 
litis study hclween 1001 and 1003 Twenty WO Itt 
originated Irom Wnktrrka (Kiues Canyon) National Park 
MMUh-weslol Alice Spurns \1\ 2()'..nd25 20'S. 130 SO' 
u» I C 43'£). 23 from an area norill of liar MncDoinietl 
Ranges horn Citen Helen Gorge to Vmnbah and Alconi:i 
Sialnnu i:?' 50' Ui 2.i 45' S *H.V JO' to i <4 SS'B) 15 
ucur Mount Davidson in the faiiaim IX-seil Wildlife 
Saiiuum), north-west \-l Aliee Springs (20" 20' lo ?0' S3 1 
S. I H? 1 25'tU 133 : 55'K), ti in die Davenport Knnge.s thai. 
Murray Downs Station via Burrow t'reek (3U s 0'|o 2V 1 

IMV s." I24 1 HI' i" 124 :v l.i. SI farm Ufa Nut* 

(AlpumdolauO (20 2o'to 21 UPS, 137 50' to US" 10' 
l.i on the Barkis Tablelands. IS horn ihe northern Rtuily 
Tablelands ( 17 5<)To tO'45'S. LU 00"io I 37 ' OO'H, ami 
l l Mt«>ni Manakat lr> KiilheniK- (12 *5'lot4 30' S l*j 
20' to I 47 " 20>;t Mmlh-casl ol Darwin, 

The siomaths til the Lais were opened so Hull uudiycsied 
contents toitkl he identified and Ihen Hie entiic g^StfO- 
micstiiut Wfltf* were preserved in 10'c formalin in 70^1 
cthanol. The preserved material was transported io Adehiide 



Tabi i I fiwtifence til hiunstintti hetmtuth fmuwftti fituntf 
nt i$K laui i an from tin Ntwtktwn lirrttow. 



Parasite, 



Prevalent [%} 



Aluindance 
(meant 



Acanihoeqihula 

ilnricolti fhtmnto.vtuim (S5 4 

Nemaluda 

Al>hrr\falti hitsl<i\pii ulo 4 i 

Ancyl&Ktonui nmhfum o s 

4w xto.stiwitt lutnu'jotvir i :,x 
( 'viiihosfinittt ttnwmufi* \. 3 
Phwnlnplt'W }>iav{>tniuiisM)A 
l'n\f» itiit rati I I) 



C'esiodd 

Sfinfmrlrti erhhit til 
Kwitiii tifcumrfnitins 



14.4 
47,-U 



I ooviiM.H 



1 10 (121 

5 
1 -3i |.h 
I 13(51 
I -51 (S» 
•i I5(0i 



i -25 (hi 
1-70(7) 



where pai.i sites wen ivnmud and WUflicO LLM1U? ;| 
di-.sn -line niii_'ntsco|U- :ind hilei'. afk'l lhi-\ hail been i'leureil 
in laeiophenol. id"etilified employing a L-i»iiipitiiiid 
tnicm-setipi.-. When present, ieci.il samples \\c\c exaitnned 
for pro(o/oa usine rrnlrdiitial lloijiuni in •^itiiuik-d MgSQ, 
sutution 

\lun of the hehtnnihs iuuml t'liible 1 1 haw beta 
icpoilcd previously in surveys nt Teral tats fi'uni New Siwilli 
Wales. Vieioriu and Tusniania [AncvfoitfuVUi itiharfi'iint; 
C\uilia\f)inirn ilusvtirntis, TbfrttUJtt ran. Sfuromeint 
VtiiUUii. Ifnni't hii-iiiatt>nrr,i\\ iiltlutUjOi llWK UIV 
.anifiianl diffeietkV.s in pivvaletiee heiween *(.nes Ihf 
principal dilfeifiki-' lu-tween iliisMirve> ami pivviotis oitcs 
i* (he very low prtvale^cc m 7 tf/ti in ihe Nordicm 
Toinoi-y Only Oil'/ ol ihe eals examined had no parasiies 
Avanlhneephalaii p;iriisiks ul llii nny in leral c il 
poptilniton?; el>cwhere have l>ecn rclerrcd l<> a>. QtyfyvfO 
sp ] '■ '. Sthiiiidt' ulemiticil the .-.peeks ,is (in, twin 
l>t>iiuth\\t'»>ti Until llv dinyn. (jiuis fnntifitins Jiukh. anil 
feral eat aei as dtliiniiye hosu. with larval Maer>. uuuurrfiig 
l.imIlt ihe skin of .i smii-ty i\t pjiSNerine hird.e 0nacftkt 
pi'/tnttoMomi was tliC mosl eommunly deieek-d pinasik- in 
ilns studs rind was also present in l.irv'e nnmhers in many of 
Ihe rats. Atllnm^h wiinnv weic luuiid with thrii pioboseis 
deeply eniheddcd into the intestinal mtiema. the assoeiated 
paiholi^ieal iL-aLiions were litni'ted to an itilkiitimai"i y 
ilUUImtP annind |lic pinhnsLis. dikvlcd ishi-n liistoloeicul 

seciittcs' svere examined. This parasite was n*u nxuveivd 
Irntn 10 eats m the uiea smith Crl Owrwin hm was present in 
animals hrun ah oi it! areas of Hie Northern Teiotoiv. Ike 
laive numbers ul O, ponuito^unni found in eats suggests 
thai birds constitute a siyiidkani item ol their diei. allliou^h 
a la:k ol .Jala on abundance Of 'bis p:ir;isHc m pauknK. 
hosts piesent i more detailed conclusions from bciny. drawn 

PIlYSiihipttTtt {IMPfiUtHilh and Ahhn\imu }uiMusf>itufti 
have been reported from feral cats and dou> | rnl n the 
Northern Territory oil a single occasion, The evidence here 
conlmns that /' f>nti{>utiniis is a iclalively comiiuin pavasUe 
nt' Icral eals m ecntial Australia. Ahhnxnaa iut>h{spiaiht i- 
nonnally a parasite <>\ varanitl liyards" hut. app.iienlly. Will 
develop also in the stomach ol cats Ryan- luuiid a 
'Vli\Mihtfthui u sp. present in cats in New Soudi WalcS bni 
ut that time dismissed »l as an incidental |>arasiie It may 
have beer T. luiMospu tiiu but Ihe lot k \t\ deposited 
j-u' aniens makes it impossible to confirm this hypothesis. 

Atiivlosunutt tuhtit'fotvu' is ^cncrallv considered to ht» 
Ihe common liuokwoiut of domestic eats' but records to 
t(aic -ugex'st thai il |* nucomiiioii in ferul cats cscept m 
iho>^ collected in the vicmity of .Sydney-. \. iiihtnfornu 
was widely distributed m the Norlhctti Icrittory, oeeinrmi: 
most iVcquently hi cats Horn the Kin^s Canyon area, the 
MacDonnell Kane.es ami Muriay Downs Station. Inlectioi; 
consisted of small numbers o( Worms (maxinium numbei - 
It) A single cal was found inJ'celed v\nh \. ivnmunh it 
parasiie moie eommonlv found in eiinids in Northern 
Ausiijdia H : this cal harboured only ftVfi adtill worms. 

An immature specimen ol (UiulltosUwui \('im\;vnini was 



170 



M.(J. O CALLAGH AN 



louiul in Ihe stomach olacal collected in KlOgK Canyon, (i, 
Kpinigerum has been found sporadically in feral eul.s in 
Australia'' ; ' but appears to be uncommon. 

The identity of Taenia tat'niat'fornris was confirmed by 
counting and measuring the large mid small loxtellar hooks 
liom 13 cestode scoleces and comparing the data with those 
provided by Verster 1 ". Spmnm-im erhmcei was detected in 
all of the cats Collected in the area to the south ol Darwin 
bul in only ciyht cats ('com the other areas ol' the Northern 
Territory. J hi*, may he due to the fact that ihe dr-.i 
intermediate nOSI Of this parasite is a fresh-walrr crustacean 
ol' the genus Cvr/ry/v 11 and the pools of fresh water 
necessary for its transmission are less, frequent in arid areas 
Parasites such as Dipxltdiuin t animtm, Ihtrinaria 
\ii/h>rrplmla and Cylicoxpinwt fcihtcus, present in other 
surveys, were no! found, 

Faecal evanimalions revealed two species of coceidia. 
haspora ((lis in 9.6$ of 1 4<-* cats and / rivnfia in O. 1 )'/ . 
Both are common parasites ol caLs and can cause disease in 
younger annuals, The majority ol the cats examined were 
'hJulis and this could explain the low prevalence of 
coccidi.in infectious (including the ahsence of 'foxttpktsntu 
y,tnuiH) in thl8 and other surveys 1 

Additional iiemauHies. I <htnomtna vmctuft), Kirtnluuit 
mt\Htir\i and \\attati\irntixxhts ch-noli were rarely found 
and (licit presence was presumably the result ot tlic 
iiI'.vmhhi ol native mammals and reptiles which are Ihe 
noiuiiil hosts ol these parasites Similarly, die ingestion M 
birds would explain the presence ^\' female nematodes ol 
ApWCta sp. In the stomachs ol two cats, Other parasites 



collected from intestinal contents and faeces but obviously 
related to the ingestion of rodent hosts were ihnnnk\\ sp, 
and P\ott ! ti>aics sp. X&tapxSilto \rxah!i\. a Ilea found 
commonly on Ratfas viliosiwimus was found in Ihe 
stomach of cats from Ihe Lake Nur>h aica and llie mile. 
/.iit/apy hapulofi, a parasite of NptOtJWH "-pp.. was found in 
cats from the Tanami Desert area. Another accidental 
parasite recovered was Sxft/tunu ulnwhiitt an o\yuiid 
parasitic in ihe caecum of rodents. 

The information presented here identifies n greater 
variety ol parasites ocCLinring in feral cats in the Northern 
Territory than previous sLudies have found. It identities /' 
firttpptiiiufa as a common parasite and demonstrates that 
cats are frequently infected with A. hasSasjnndir h also 
confirms ihal the feral cal pteys on native mammals, birds 
and icpliles. This is particularly so in the case ot birds, with 
ihe Initio numbers rrf i) pifnnila\h>mi Suggesting frequent 
predalrofc 

This work was begun by Murray Barton, when employed 
at the Arid Zone Research lusliuuc. Alice Springs and was 
conducied on material collected by the siall ol ihe 
Conservation Commission oi the Northern Tern Lory. Alice 
Springs We wish lo thank Mr Ikirton and all ol tin 1 
individual olfjeers who kindly collected and preserved the 
viscera M the cats ami David (iilv.on for his help with the 
manuscript 

Representative specimens o\' all of the helminths 
collected have been deposited with (he Australian 
Hemmiliologica! Collection, South Australian Museum. 
Registration numbers AIIC 20181 to 31)231 



'Jones, R. (1983) 'lernl Cat p 4N9 hi Strahan. U. (lid.. 
Complete Bonk of Australian Mammals" (Angus & 
Robertson, Australia). 

Kyan, (.;. K. (1976) AusL Vet. J. 52. 2:4-2:7 

Ionian. B. .1. 1 1972) Ihhi 48. 133-136. 

Gregory, G. G. & Munday, B. U (1076) U>iJ. 52. 317-320. 

Cnman, K. J„ Jones, E, II. & Driesen. M. A. ( I 98 1 1 Ihal. 
57. 32 4-327. 

Schmidt, G. D. < I9S3) J. Para.silol. A'J( 2). 397 399. 

Barton, M. A. & McRwan, IX R. (1993) An i Vel J, 
70(7). 270. 



Junes, H. I. ( I9S3) Aust. J. /.ov\. 31, 2S5 :9X. 

PrescotI, C\ \V. (19X4) University Of Sydney, 
Postgraduate Koundation in Veterinary Science, Review No 
24. Parasitic Diseases o\' the Cat in Aust. 

"'Verster, A. 1 1%0) OndcMcpoorl J. Vet. Res. 36( I I. k5B. 
rhmsmore, J. 1). & Shaw, S. K. (1990) IJnhersily of 
Sydney. Postgraduate Koundation in Veterinary Science. 
Review No. 31. Clinical Parasitology of Dogs. 
<fievcridg«, I., Presidente, P. J. A. & Arundel, J, H, 
(1978) Aust. Vet J. 54, 46. 



M. G. O'CAI.LAGHAN, Prinutry industries (South Australia). GPO Box 1671 Adelaide S. Ausl. 5001 iuu\ I. 
BEVERIDGE, The University of Melbourne. Veterinary Clinical Centre Princes Highway Werriboe Vic ; M)M). 



PREDATION OF EARTHWORMS BY THE LAND PLANARIAN, 

AUSTRALOPLANA SANGUINEA (MOSELEY) VAR. ALBA 
(DENDY) SENSU JONES, 1981 (TRICLADIDA: GEOPLANIDAE) 



Brief Communication 



Summary 

Earthworms can significantly reduce soil degradation and improve pasture and crop 
production 1 . Land planarians (terrestrial flatworms) can, however, greatly reduce the 
abundance of earthworms in agricultural fields. For example, a New Zealand 
planarian, Artioposthia triangulata (Dendy), has been accidentally introduced into the 
British Isles where it now feeds voraciously on local earthworm populations. The 
implications for crop and pasture yields and the abundance of wildlife that feed on 
earthworms in the British Isles, are of major concern 2,3,4 . Another planarian, 
Australoplana sanguinea (Moseley) var. alba (Dendy) sensu Jones, 1981, has also 
been accidentally introduced to the British Isles, either from Australia or New 
Zealand 2,5,6 . Relatively little is known of its biology, in particular its feeding habits, 
except that it will feed on "earthworms". 



(twf\iuiu»>*i'1 iht fofatfatiiNfiif&AHHAWtt, I20tlr. W-llH 

BRIEF COMMUNICATION 

PREDAJ ION OF EARTHWORMS BY THE LAND PLANARIAN, WS7RA1 MP1ANA 

SANGUINEA (VIOSEEEY) VAR. ALBA (DENDYt S7L\ r SV JONES. 1W1 

(TRJCLADIDA: GEOPEANIDAE) 



I-aiihwnum can significantly mduee mh! degradation and 
impiovc pasture ;inil caip production 1 Land plunnrinns 
(terrestrial llatworms) can, however, ^really reduce lite 
abundance of earthworms in agricultural fields. Fflr 
example, a New Zealand plaiiurian. .lr//o/v,',w//m 
hutuvMhiht (lAnitlyi, has been accidentally introduced into 
iliv- Kuirsh Isles where ii now feeds Voraciously on local 
carihwonn populations. The implications Tur eiop and 
!',i' ihi;' yield:- ami Ihe abundance i»f wildlife lital iW'il nn 
railhwoi ins in the Unlisb lsk - s. ale tit major concern . 
AnoihcTplanariau. AustralnftliitutMtnuuiniyi (Moselcv i vur. 
ii(h>> <l>endy) sviistt Jones, l%l, has also been ;tochlenl.iIlv 
mtn'duccd l«> Hie RnlisM Isles, eilhci foul* Auslraiia or VfW 
AaluiiJ' ' '. Kelalivch Utile is known of its biology. Ifl 
paiheiilai its leedme, habits, evcept ihat M will feed on 
'oaUhwrnim.'' 

\usiiulo{>Uihu sprues are widespread and Iretjuenily 
eueouiilercd in man- modi lied habitats in eastern 
AiHralia*'. Recently, .-V MtMjitimv vnt~ •tlhi \u> Ik-cii 
round (0 b*5 nctihei /J. .uti>\:nwtui nor A. ulb<i but in in tail a 
ito\s pcnus and species il Wiusnr. pcis coimn ) This 
Hniwonn s only Known occurrence in Ausitali'i is i[\ 
Kime-swood, South Australia. alUit>u^h il probably occurs in 
other urban and disturbed liahuats. 

Hit 1 liflnciil surface ol A. WiXWfiUM var, ,t!h>i u niosllv a 
uniform (m\t .'ii«init-h*MWii colour with a decpci brown 
peach anterior up: tis ventral surface is white \ \ttny.nincn 
Vttr, itlbti v\as commonly found by one &T us (T t; .Tj in 
moist habitats, such as under bricks, rocks and piccc< ol 
wood and. sometimes in the top 10 an ol soil inn suhuihiin 
eajden in kiueswood li ted on garden earthworms when 
ollcrcd them tu jilitslrt. j.u 1 -. but mil on oll'ici' uiVortobottL-s 
such as J. noils, -.lu^s. miihpcdcs. slalors or c'arwign. 

Tlnn impef docunicnis five earihworni .species upon 
which A, htjwtjfltfi v;tr tiltm wdl Iced nndi-T !;>♦*»* n;it» n\ 
civnditionN and pruvidc\ some da(^ fffl its reproduction .ind 
early developuicrn The w«rk loruis pan of a broader Siutl} 
jimed at itTipri.»v\d itmn;i^cmenl of earl]iwiMO*s jv a 
rcsouivc in snuih LM\u-rn Ausiralia'. 

I irsily. mx adult A, \wi&utn<t var. itlhn lappros. 0-.S. ;ni 
I.Mhj) \u-tv n-HctiL'd Troiu the Kin^s\w«od garden in 
(Xnibcr lUO-i These pijnmintis v\ere placed in pans in 
three opaque, closed plaslir i.<«ntaint*rs (duin. 9 cm; ftcVil! 
IU cm) and mtu'niaiiied ul 15 C- The hoikmi »f (audi 
eonlainer was DtvVcrtcJ wuh 1,5 cm of^iitlen soil which 
was kepi moisl hy spmymy rctrulaily w'nh wak*r. i"i\e 
c\«aiL i-ariliworm s|»ccies. AforrfCfotloa aihyihosn 
(S.iviy.nv i. A, mh'<i (Siivi^iyi, A. ntynj&tdffl tDtipes) ajid 
.\ itmy.tt iWd^-i tlAiinhiicidJcl a^<\ Afiri^:\coU\ tlttbhts 
1 1 led lu'i > iAcanlht»dnhdae) wert* nllcied In the planai urns 
•is l»Hid. Theie ea'th^ocm species, apart Irum A, hfy}$t* 
which (•• widespread ifl and was obtained from northern 
Tasmaiu.i, aic commoniy louud in ^irdtns and .u^rieulmrat 
fiolds m Si »uth AtiHlralia and Vicloi la " ; \\\o inilivitltlals o) 
.1 pailicular earthworm species were introduced into each 



L'oni.umi wilh tltr llatwurms. mte earthwonn hflUuTUf 
at a lime. TredvUion was aw>sed OMfl » pgji(»U «>f ?| (J- 
BaUhw4>oii sptatLs wen. L-kme_ed in the cv-nlaincrs Mihctl 
predaiion <»ccmtcd -Ml speeii-s of earthworm were killed 
and rau-n by A. KtiftfflltRfi vai. wflbfl, The planarians a|.* df 
rtf p.iO Ol an eaiilnwnni. All pttHS '.'I die hodv Were 
anacKed namely head, tail and mk!s<xiioii- In sntne KEDtoti 
\Ahere onl\ tlw tail wa-- eaten. Itft earthworm simww^i. in 
eonlrasi, eaitlisvoiui^ ahsJVs dn vVKen parlial|' f 4 -lien hj V 

tritt'!;*Uhltu 

Secondly, seven aJnli \ ifWttmUvUi v ar. dhu, RtW3 ife 
same Mnevwtitid gyf^fi^i *t'tf placed til sepaiatc pla-li. 
cxntiiincrs. as Jcvjihcd ahose Two ailull \pnnvri,nU'ti 
,vht'Wi'\it wea' added til kSOiHl i-oniamer. rhe mimhets o\ 
mi.'^ine or patoalK ealen eatihwonu 1 veie receded 
appiuMliiatcly every 2d li ft* 3d d Larthworms Wiav 
teplaeeil if eoniplele <>r partial ptcdnlton had ouvurred I v," 
plWftirm05 died durin? [llC ( -vpci-im^ni , l-nfli al Aa\ 2S. Uu 
avci.-eA earthwonn.-v were attacked once cveiv s d pfli 
|>lauarian iranev j - I j j|j. This is siimlai lo the I'aic at 
whuh A. >:~>ii!ivn!(tt<t altaekcd aiK>ttier eatlliwonn sjii-nes, 
Inittiti fr/'(/(MSa\ien>M ].4alUck> per plananau per week 
at 10 C) : . 

(Jurine the UN cvperiment. the sn planarians ileposiied 
live eiti>-e apsides between them. In the second experiment, 
the ^even planwnans tlepositeit 17 ee,y cap.sulcs (4 40. I. 
2. 3 and 3 each), tlie cnpsule^- were sphcraal h> oval in 
slun»e and un^ed horn J - 4 mm in diameter They were 
maroon red in colour and were deposited cither on the soil 
surface or a lew mm betow it bui were not attached to the 
soil surface by stimc as \\.\\ been noted Ivy terrace & 
Baker lV»r C>u/tot>l(tiui uaviilfti lMosclc>). The c^e 
capsul-s were maiumiiied at I 5"C Ul opaque, closed plastic 
conlaincrs (>Jiam, 7 cm, hc^ht 5 em) hti Up tu I4 U d 
i uurieeii ol the capsules hatched. These successful capsules 
were deposited belweeil 3 1 Oclohei and 1 3 Oeccmber I l 'VM 
and all hatched during Ee&rtWfii r^t^ t tytty hatchine imie 
w-j-- S7 d [range 71 11)1 d), Forty one luvenile planaiians 
wcie rearetl from die eap.sule\. I - 5 from each capsule 
intean — :, J- i. r pon entcrgenec, lhe.se juveniles were pale 
vcllow in colour wilh a reddish anterior Up. The nuiin tntdy 
colour darkened to eilhei a givv or pale orange - hmw n ovei 
a 3 It ii period. The juvenile planarians were provided 
with iuvemle A, rUtfginv&Q as food Uul fib.) and beean 
leedme on them after an avera^tf of 10 d panee h 22 d). 
hiiihl tjf Ine juvenile plunarians reached uduli %\f& three 
inoiiihs after emergence. These eiyht indn'iduals were dien 
erouped i-i one container, and, after a lui'thef month, 
produced 1hree bgg capsules beiwcen Ihem. 

Introduced lumbrieid earthworms are generallv louch 
mote abuiiilani than native species in urban and agricultural 
soils m southern Australia' 1 ' Native eniilnvorms are 
presumed (o be more abundant in uiidi.slmhcd habitats, 
where invasion by iniroduccd specie^ iv believed to be 
limned 1 - 1 "\ Whethei or noi ,\ KtrnjWfffrrti Vji ttfba prr>s 



178 



upon native earthworm species or significantly influences 
the abundance of cither introduced or native species in 
disturbed or undisturbed sites in the field has yet to be 
determined. However, given the numbers of earthworms 
eaten in the laboratory experiments ( I worm / planarian / 5 



d), the influence of A. sangumea on earthworm abundance 
could be significant. 

We thank Don Terrace for help with the collection of the 
planarians and Leigh Winsor for assistance with 
identifications. 



l Lee, K. E. (1985) "Earthworms. Their Ecology and 
Relationships with Soils and Land Use" (Academic Press, 
Sydney ). 

Blackshavv, R. P. & Stewart, V. I. (1992) Agric. Zool. 
Rev. 5, 201-219. 
'Blackshaw, R. P. ( 1995) Acta Zool. Fenniea 1%, 107-1 10. 

Boag, B., Yeates, G. W., Johns, P„ M., Neilson, R., 
Palmer, L. F. & ligg, R. K. ( 1995) Ibid. 196, 212-214. 
'Jones, H. D. ( 198 1 ) J. Nat. Hisl. 15, 837-843. 
''Anderson, R. (1986) lr. Nat. J. 22, 141 146. 

Winsor, L. (1977) Proc. R. Soc. Vic. 89, 137-146. 
Winsor, L. (1979) Vic. Nat. 96. 155-161. 

Baker, G, H. ( 1 996) The ecology, management and benefits of 



earthworms in agricultural soils, with particular reference to 
southern Australia /// Edwards. C.A. (Ed.). "Earthworm Ecology" 
(Soil mid Water Conservation Society, Ankeny). In Press. 
'Baker, G. H., Thumlert, T. A., Meisel, L. S., Carter, P. J. 
& Kilpin, G. P. (1996) Soil Biol. Biochem. In Press. 
"Terrace, T. E. & Baker, G. H. (1994) J. Anst. Ent. Soc. 
33,371-372. 

1 Wood,T. (i. (1974) J. Anim. Ecol. 43, 87-106. 
'Abbott, I. (1985) Ausl. .1. Soil Res. 23. 263-270. 
"Lawson, L. M. (1993) "The distribution and abundance 
of native and introduced earthworms in an area of pasture 
and native vegetation near Cape Jervis, South Australia". 
PhD Thesis, Flinders University ol South Australia. Unpub. 



T. E. TERRACE, Division of Soils 
S. Aust. 5064. 



nd G. H. BAKER, Division of Entomology, CSIRO PMB 2 Glen Osmond 



MASS FROG MORTALITY AT TWO LOCALITIES 
IN SOUTH AUSTRALIA 



Brief Communication 



Summary 

In recent years much attention has been given to the alarming decline in amphibian 
populations worldwide. Australia is no exception to this with severe reductions in 
population size recorded for several species 1,2,3 . Suggested reasons for apparent 
declines vary widely, with habitat alteration and fragmentation, acid rain, introduction 
of exotic animals, toxicants, increased UV and pathogens being implicated in several 
instances 4,5,6 . It is possible that such factors also act in concert 5 . For example, habitat 
alteration may sufficiently stress the frogs so that their immune systems become 
compromised, making them more susceptible to infection 7 . It has also been suggested 
that such declines may only be a small part of a naturally occurring cycle of 
population flux 8 . 



hmVH Htm ttfthfi tywj tin fat Hi S. ilfiWi \ IMft, I20( 11 I fl 



BRIEF COMMUNICATION 

MASS I RO(i MORTALITY Al TWO LOCALITIES IN SOI' TH Al'S TRAUA 



in .cll-iii vcjis iviui.it aik-nimn lu-i been gbfti w Ihe 
alanmuc deUliltO ill amphibian pnpulaiions worldwide, 
Ausimha is no ewcption Id rhJj, uiih seveie reduclions m 
population Kt/f reeoidrd lor .several speiacs - '. Sityar.-aed 
reports, lor apparent declines vary widely, widi hubiiai 
alleraliou ami lra\:iue.nlalion. uptfl ruin introduction ol 
CXULH* MOtnUill. kft'tVUQt&i inrrviisn! l.V ami pnlhoyens 
Iilmmj implicated in several instances^" II is possible ihui 
such faetois also ael in eoneen : '_ For example, habitat 
alkralion may sulTicienlly s*ivs^> the Crops w thai Ibvtt 
immune -ysk-nis become compromised, making idem more 
susceptible 10 inlt.Llii>n\ ll litis al-.o h^tfli oiL*i?csk'd that 
Mich declines may only be ft iJtfJ&ll PWW "'" ;l n;t|ii):<U> 
huCurffiijrfycte olpopulaiion lluv 

lit ilk* Held Ihe ohserv/nion Hf MiVjyrBl fefciul Ini^s 
Miimhuite-oitsly |5 unusual as ihey degrade quickly alter 
denlle Hence the piestaiLe til multiple ljiqivm". i-- j si 

mass mortality til a sJuw time (runic iFiu ftugs thK would 
probably be |y,\* lliail llffCC itfljjs) There have been ten ill 
icporls "I such :i phenomenon fmrn soverat tey.mns 
including Swii/e-rlund". Hawaii ° I upland and Australia 11 . 
Heie WW report max., (toy umiijlniex ,ii tVYO rDEUlRlcS in 
Noulh Auxli alia. 

I lie MM iiMikni look place mi 2") May Ifi&l Rain al 
Nmutolpa. .South Australia 'latitude _U 2M' S. longitude 
|39' "' p) filled a dam on :i vineyard (n a depth ill 
appio\imakl> (Xl5 hi. Nuinciou.s burrow m«_* friigl 
[Nmt/ntiMhito flfiMtai emerged ami. on the (ollowin^ tia\. 
many were found Jc*lJ ;<l 'he ede.es of ihe .lain Ope i.il us 
Al I I I veiled the site on -M May [992 and ivcow tm ! ' 
dead specimens. All were :u ihe edge oldie ualei wliliiH 
had atreaily receded [0 .t dcplli ol' f)..i ni ^oini hail ludtl 
sp.iwn hill oilier dead. uiaVid mdi\nlllals were lueakd, The 
spavMi had dried Dill and il could mu be dek'rminc'l il ihe 
DVil llUlI been leriilKed. 

Ihe Mmidlaueoas nahliv ftl" the dealh ol Ilk- Iiols 
iiidiealctl llic presence nl a pOlllltflTll. Accordinj.' lo ihe 
Ludowueis The onl; ( chemical n>ed al ihe ^ik- was >pol 
%prayinv ol ll'e herbiLidc Roundup I inanLdaeLUied hv 
A'UmMinlo aid consiMine ol glypliosaic and a dispet'sanU lo 
conn ol ihlMlcft. Spi'ayiii" had been itinleri.iken 
appniMMialelv Ihree iitoillliv pte\H>Li;^ly (in IVhiuary l^'-'Zi 
This ptnod shtuild have been adequate liu ihe de^radalmn 
ol Mil hed'ieide 

'Tyler, M. .1 I |WyJJ Alvie,. % -M 5d. 

Kit hards, S. .1.. McDonald, K. K. & Allord, K. A. i |*Wa i 
I'm'dK (un ■.ecvaiion ISioloiiy I. (»o-77. 

yiftl, J. L, & Savlin; I,. ETWl Ihe sifiuis ol amphibian 
populations: a eompdalion and analysis WnrkinL* 
doiununi i IUrN/&'SChfltWittllgAwnlnhiiUlTllitl' Fiticc. 
VVoild ( oiiseivalioil lilion. 
AVyman, K. L. I IW1| ( tunservnlioii Hiolouv 4 =15(>^5I. 

cinxy, C« vS. Ury;im, C J. I W5) BttVii^nrncrtltil ttcaHli 

pi r.peelive 1 - 10^ Suppl. J. I ^-1 7 
lainrniiee. \V. !•.. Mt Dmiald. K .K., & Spcare, U 

( 'oiim'iv'iirioii rtidiouv hi Ptqmti 
Carey. Ci 1 1003 ) te 7 555 3fi5 



The second sile ol ttliiSS morlaliiy was al I'ualana llol 
.Springs, 27 km nonh-e.isl ol Arkamoln ||oiiies|e;i(l m llie 
nonheni Flinders Ranges (I'M W?oDfl on Yudiiamnlana 
1^7 ^T- 1 1:51)000 lopoui.iphie iiuipt Vhi- -. K a ^eolheiiually 
heaied body ol walci llirou^h which lailon ^;i^ babbles. 
W.iief emei'L'es al 60' C and has a radiikiciiviiy \<( 
approvimalely 20011 pPaf\ Allei sulliLien! landall IVesh 
waiet can also be loimd upsiicam ol' thd hot spring. On ^ 
Decemhet 1^4 nine <Scih\ fiogs were disco\ei*ed b> C'.W. 
rio;iliiH- al Ihe ho| sourer ol l|ir -prm^ :aid. h\ llieir sliile. 
weiL -.oiisidcrcd W have )\cc\\ dead lor less than two davs 
Seven | intludiliL' one niveinlei v,eie the spotled yrass |'p>si, 
{.i>itnin/yiht.\ns Uisnmnu i/v/i. .md lln.- olhei I wo weie 

lepivseniat. *»..-. »M the red live Frog, l.itoriu rtihtlht Iheir 
advanced Male o! deeay piecluded ii iiieaiMnplul 
inse'di^aftrn by palholopi-.lv. 

Despite exlensive SoHtvTlM-, Ho liVlfig y.^////f»(/\7/th/cv 
hiMiitiitiitt.sis were di.srmeied :il iffc MlC \Uieteas iiuu'm 
itthctJH was heard calling but no| seen. In cooler parts ol (he 
spi'iiic drtvUttStr^ffltl. there vvl*iv holh (.aval -md ailnll 
luteieis it rutin nfuniin and 111] dead Irons wcie loniiil 
Tlieie \>b->.rxi.iioii- were made die day alter 'he lll'st 
oeeuMon lliat oiin hiid kdlen tor one monlli. 

Mass moiialihes were Ugatll rccoidcd on 8 May Mt%RI 
ihe sumo she, I leven vlead C, lipntiii were found bolh al Ihe 
hol source and ap-aa-aii) oT il in iMilakd |)o*tls oleouUi 
Wilier i2V It. These ineluded bolh adulls ;iud pivemles. 
Several livne kuval. juvenile and adult C. riparitr and /. 
knin/unriisi\ were also observed al the mIc mi (his dale. 
Occasioiiid sigllli^S al dead frogs ikmlini! in Ihe hO( pU0l« 
ba\e been made reeulaily nvcr Hie lost ?{( yuars tl). Sprite 
per-. c'Hiini.K 

The presence of such a I&rgti nunibci ol dead I'ro^s is 
su^i'esijve of a paiho^cnic eiuisC- Tin:-, oeeiinvuce eannoi 
be al I ri hiked readily lo any lonii of anlhiopo^cnic 
iiueirefeiiee. a\k\ while h.lgtl \^ak'|• lemperainre ma\ have 
eonliibiih'il lo llu- m;iss inoHiihly. il is uuhkelv lo lime been 
Ihe single cause o| so inauv deallis v, Ulun ^uch a %hori lime 
Although tavesligaiion into amphibian palhoiieiis i> snll m 
ils inl'ancy they have been previously linked wnh lm^ 
moriahties Unh iti Australia'' ' and abi'tad 1 

Simon Sieuiboiivr Adnau liradlord. Meven Walker and 
Kebeeea Shori nre ih;mked lor dieu .issisiance hi die held. 



PtihuuiiHi, J. II. K., Seott, I). K., Senditsih. K. O., 
Ottllwelli J. F« Vilr, K, .1. & WhilMd ( ;ihlHins J. ( | L >9 1 » 

Science 253. S02-805, 

'Tnmloji (l l '"2' New-ktier ol die U A N,\S( la-k T>rcc 
dm Declining Amphibians. June l^ L| 7 (?) 
(, <ireen. D.K. i loo4t ke|>ori produced Tor llic Disease !ffld 
Paiholov>y Working uionp. Declining Amphibian 

Populations Tjskloiev 

Dniry, S. E. N\ (oiuyh, H. K. X. CuMiiiiiyliHni, A. A. 
(I^Jfii'Vek'tinaiv Record 137, 7?-7V 

iullen, B» R. a Owens. 1 . & \\ 'hilliiiRitm, U, Ji I 1^951 

n '-><. WC5 M| Auiflttli 'Ar^anisms 2.V S.^-o;', 



VI l( IIAI I J TV I TKamiCKAIf; K WII I IAMS. Do pari men I olZooloin. (i|iivt'i>ily of Adt'lauL S Ausi, 

sow. 



ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED 



Patron: 

HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA 
SIR ERIC NEAL, AC, CVO 



OFFICERS FOR 1996-97 



President: 
T C. R. WHITE, BSc, BSc(For), PhD 



Vice-Presidents: 

M. DAVIES, BSc, MSc, PhD 

M. A. J. WILLIAMS, BA(Hons), MA, PhD, DSc 



Secretary: 
O. W. WIEBKIN, BSc, PhD 



Treasurer: 
J. H. BRADBURY, BSc, MSc 



Editor: 
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Assistant Editor: 
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Librarian: 

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GradDip (Inf Studies) 



Programme Secretary: 
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Minute Secretary. 



Membership Secretary. 
G. A. CROOK, BSc 



Members of Council: 

P. KOLESIK, BSc, PhD S. BARKER, BSc (Hons), PhD 

N. A. LOCKET, MA, BM, BCh, PhD, DO M. J. WRIGHT, RDA 

R. D. SHARRAD, BSc(Hons), PhD, DipT(Sec) 



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