VOL. 111,
29 MAY, 1
Contents
PARTS | & 2
987
Transactions of the
Royal Society of South
Australia
Incorporated
Tester, M., Paton, D. C, Reid, N. & Lange, R. T. Seed dispersal by birds and densities
of shrubs under trees in arid South Australia - -
Beecroft, A. S., Cann, J. H. & Stocksiek, C. A proposed reference section for the Tortachilla
Limestone -
McKenzie, K. G. & Guha, D. K. A comparative analysis of Hoceue/Obaotene Sounslary
Ostracoda from southeastern Australia and India with respect to their
usefulness as indicators of petroleum potential - - - ~ -
Southcott, R. V. The classification of the mite families Trombellidae and Johnstonianidae
and related groups, with the description of a new larva (Acarina:
Trombellidae: Nothrotrombidium) from North America - - -
Southcott, R. V. A new larval mite (Acarina: Trombidioidea) ectoparasitic on an Australian
centipede, and the Trombidiidae reclassified - - = - -
Thomas, D. P. New freshwater diatom taxa from tropical northern Australia - -
Arumugam, P. T. & Geddes, M. C. Feeding and growth of golden perch larvae and iy
(Macquaria ambigua Richardson) - - - . -
Birks, P. R. & Olsen, A. M. Pesticide concentrations in some South Australian birds ae
other fauna - - -
Beveridge, I. Echinocephalus overstreeti Deardorf f & Ko, 1983 (Nemnstone:
Gnathostomatoidea) from elasmobranchs and molluscs in South
Australia - ~ - - - - - - - -
Houston, T. F. Fossil brood cells of stenotridid bees baer eben: Apoidea) from the
Pleistocene of South Australia - - - > - - -
Lee, D. C. & Pajak, G. A. Anoplozetes, a new genus of Zetomotrichidae ee as
Cryptostigmata) from South Australia - -
Gowlett-Holmes, K. L. The suborder Choriplacina Starobogatov & Sirenko, 1975 with
a redescription of Choriplax grayi (H. Adams & Angas, 1864)
(Mollusca: Polyplacophora) - - - -
Gowlett-Holmes, K. L. & Zeidler, W. A new species of Reahthoehitena (Mollusca:
Polyplacephora: Acanthochitonidae), from South Australia - -
Brief Communications:
Peterson, M. & Shea, G. M. Reidentification of Ctenotus schomburgkii (Peters, 1864)
(Lacertilia: Scincidae) - - - - - - - - -
Storr, G. M. On the identification of Clenotus schomburgkii (Peters) -
Angel, L. M. & Mutze, G. J. On the occurrence of Brachylaima sp. Cirematoday in he
feral house mouse, Mus musculus, in South Australia - - - -
Johnston, G. R. Reproduction and growth in captive death adders et tel antarcticus
(Squamata: Elapidae) - ~ - - -
Pring, A., Snow, M. R. & Tiekink, E. R. T. Bavdlacwmnite fon South ‘Anetealis - -
Smales, L, Parasites of the wombat Vombatus ursinus from the Gippsland region, Victoria
Southcott, R. V. & Glover, C. J. M. The occurrence of Desmonema gaudichaudi (Lesson)
(Scyphozoa, Semaeostomeae) in South Australian waters with records
of fish-jellyfish symbioses ~ - - - - - - -
105
115
119
121
123
129
PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
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TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
TRANSACTIONS OF THE
ROYAL SOCIETY OF SOUTH AUSTRALIA ING
CONTENTS, VOL, 111, 1987
PARTS | & 2, 29 May
Tester, M., Paton, D, C, Reid, N. & Lange, R. T. Seed dispersal by birds and densities
of shrubs under trees in arid South Australia - -
Beecrolt, A. §., Cann, J. H. & Stocksiek, C A proposed reference section for the Tortachilla
Limestone -
McKenzie, K. G. & Guha, D, K. A comparative analysis of Enceosioligecené Rourdare
Ostracoda from southeastern Australia and India with respect to their
usefulness as indicators of petroleum potential - - - - -
Southcott, R. Y, The classification of the mite families Trombellidae and Johnstonianidae
and related groups, With the description of a new larva (Acarina:
Trombellidae: Nothrotrombidium) from North America - - -
Southeott, R, V. A new larval mite (Acarina: Trombidividea) ectoparasitic on an Australian
centipede, and the Trombidiidae reclassified - - - - -
Thomas, D, P, New freshwater diatom taxa from tropical northern Australia - - -
Arumugam, P. T, & Geddes, M. C. Feeding and growth of golden pen larvae and Iry
(Macquaria ambigua Richardson) - - - -
Rirks, P. R, & Olsen, A. M. Pesticide concentrations in same South Ausialion birds ris
other fauna - - - - - - - - - - -
Beveridge, 1. Echinocephalus overstreeti Deardorff & Ko, 1983 (Nematoda:
Gnathostomatoidea) from elasmobranchs and molluscs in South
Australia = - - - - - - - -
Houston, T. F. Fossil brood cells of stenotridid bees (Hymenoptera Paoieny trom the
Pleistocene of South Australia = - - -
Lee, D. C. & Pajak, G. A, Anop/ozetes, a new genus of Zeromntrichtdne (Acarida
Cryptostigmata) from South Australia - - ~
Gowlett-Holmes, K, L, The suborder Choriplacina Starobogatov & Sirenko, 1975 with
a redescription of Choriplax grayi (H. Adams & Angas, #864),
(Mollusca: Polyplacophora) - - - .
Gowlett-Holmes, K. L. & Zeidler, W. A new species of Avmiitochizona: (Mollusca:
Polyplacophora: Acanthochitonidae), from South Australia - -
Brief Communications:
Peterson, M, & Shea, G. M. Reidentification of Crenotus ithe PeseEs, 1864)
(Lacertilia: Scincidae) - - - - - - .
Storr, G. M. On the identification of Crenotus schomburgkti (Peters) - -
Angel, L. M. & Mutze, G, J, On the occurrence of Brachylaima sp, (Trematoda) in ihe
feral house mouse, Mis niuscu/us, in South Australia - - - -
Johnston, G, RB, Reproduction and growth in captive death adders EAM EES antarcticus
(Squamata;: Elapidae) - - - -
Pring, A., Snow, M, R. & Tiekink, E. R. T. Paratacarilte from South “aniealia - .
Smatles, L Parasites of the wombat Voribatus ursinus from the Gippsland region, Victoria
Southeoit, R. ¥. & Glover, CJ, M, The occurrence of Desrnonema gaudichaudi (Lesson)
(Seyphozoa, Semaeostomeae) in South Australian waters with records
of fish-jellyfish symbioses = - - - - - - - -
105
115
119
12)
123
127
129
131
PARTS 3 & 4, 30 November
Barker, S. Eighteen new species of Stigmiodera (Castiarina) (Coleoptera: Buprestidae) -
Beveridge, I. & Sakanari, J. A. Lacistorhynchus dollfusi sp. nov. (Cestoda:
Trypanorhyncha) in elasmobranch fishes from Australian and North
American coastal waters - - - - - - - - -
Christophel, D. C & Greenwood, D. R. A megafossil flora from the Eocene of Golden
Grove, South Australia - - - - - - - - -
Beveridge, I, & Campbell, R. A. Trimracracanthus gen. nov. (Cestoda: Trypanorhyncha:
Eutetrarhynchidae), with redescriptions of 7. aetobatidis (Robinson,
1959) comb. nov. and 7: binuncus (Linton, 1909) comb. nov. - -
Geddes, M, C. Changes in salinity and in the distribution of macrophytes, macrobenthos
and fish in the Coorong Lagoons, South Australia, following a period
of River Murray flow - - - - - - = re =
Dulhunty, J. A. Salina bed instability and geodetic studies at Lake Eyre, South Australia -
Campbell, R. A. & Beveridge, I. Floriceps minacanthus sp. nov. (Cestoda: Trypanorhyacha)
from Australian fishes - - - - -
Campbell, R. A. & Beveridge, l. Hornelliella macropora (Shipley & Hornell, 1906) comb.
nov. (Cestoda: Trypanorhyncha) from Australian elasmobranch fishes
and a re-assessment of the family Hornelliellidae - - - -
Brief Communications:
Davies, M., Watson, G. F. & Miller, C A. New records of Uperoleia (Anura: Lepto-
dactylidae) from Western Australia with supplementary osteological
data on Uperoleia micromeles - - - - - - ~ -
Reay, F. Australian plant nematodes: Longidorus Micoletzky, 1922 and Paralongidorus
Siddiqi, Hooper & Khan, 1963 (Nematoda: Dorylaimida) — - -
Sokol, A. Yabbies at Dalhousie Springs, northern South Australia: morphological evidence
for long term isolation - 5 - : - - - - -
Alley, N. F, Middle Eocene age of the megafossil flora at Golden Grove, South Australia:
preliminary report, and comparison with the Maslin Bay flora - -
Errata:
Smales, L. Parasites of the Wombat Vombatus ursinus from The Gippsland Region,
Victoria - - - - - = - . - - -
taserr te TRinsucrinns af the Royal Sociery of South Avstratia, Val il, Parts ¢ & 4, 28 November, 1987
133
147
155
163
173
183
189
195
201
203
207
SEED DISPERSAL BY BIRDS AND DENSITIES OF SHRUBS UNDER
TREES IN ARID SOUTH AUSTRALIA
BY MARK TESTER*, D. C. PATON?, NICK REID® & R. T. LANGE*®
Summary
The frequencies of nine species of shrub under Acacia papyrocarpa trees, under Myoporurn
platycarpum trees and in the spaces between trees were measured in a South Australian arid zone
woodland. All nine species were at least as common under the trees as they were in the open, and
five species had significantly higher frequencies under the trees. Three species with higher
frequencies under trees, Chenopodiurn gaudichaudianum, Enchylaena tomentosa and Rhagodia
spinescens, produce fleshy fruits that are consumed by birds. Their seeds would tend to be
disseminated under trees in which the birds perch. Enchylaena tomentosa was equally abundant
under both three species, but R. spinescens and C. gaudichaudianum were more abundant under
A. papyrocarpa than under M. platycarpurn. Traps set under the canopies of the two tree species
collected similar though highly variable numbers of seeds. This suggests that rates of seed
germination or seedling survival for R. spinescens and C. gaudichaudianum are higher under
A. papyrocarpa than M. platycarpum. The distribution of these two shrubs cannot be explained
solely by the pattern of seed dispersal by birds. For Enchylaena tomentosa, seed dispersal by birds
may provide an adequate explanation for its distribution.
KEY WORDS: Chenopodiaceae, seed dispersal, plant distribution, birds, fleshy fruits.
SEED DISPERSAL BY BIRDS AND DENSITIES OF SHRUBS UNDER TREES IN ARID
SQUTH AUSTRALIA
by MARK TESTER*!, D, C, Paton], Nick REID? & R, T. LANGE*
Summary
Tester, M., Paton, &.C., Reo N. & Lanor, RT. (1987) Seed dispeesal hy birds nnd densitics at shrubs
under trees in afid South Australia. Trams. &. Sac. 8. Aust. 114(1},1-5, 29 May, 1987,
The frequencies of nine species af shrub under Acacia papyrocarpa trees, under Myoporam plutycurpam
trees and in the spaces between trees wete measured in 4 South Australian arid zone woodland, All nine
species were at least as common under the Lrees as they were in the open, wrd five species had significantly
higher frequencies utider the trees. Three species with higher frequencies under trees, Chenopodium
gondichaudianum, Euchylaena tomentosa and Rhagodia spinescens, produce fleshy fruits that are consumed
by birds. Their seeds would tend to be disseminated under trees in-which the birds perch. Enckv/aena
tomentosa was equally abundant under both three species, hut R_ spiriescens and C. gaudichaudianure
were more abundant under 4. papyrocarpa than under M, plutycarpum. Traps set under the canopies of
the two tree species collected similar though highly variable numbers of seeds. This suggests that rates
of seed germination or seedling survival for XR. spinescens and C. gaydichaudianum are higher under A,
papyrocarpa than M. piatycarpum. The distribution of these two shrubs cannot be explained solely by
the pattern of seed dispersal by birds, For Enchwaena tomentasa, seed dispersal by birds may provide an
adequate cxplanation tor its distribution.
Koy Worps: Chenopodiaceae, seed dispersal, plane distribution, birds, fleshy frnits.
Introduction
In the Australian arid zone, a variety of shrubs
("berry chenopods") and small trees (Acacic,
Exocarpus, Heterodendrum) produce brightly-
coloured fleshy fruits or arils that are consumed by
binds (Davidson & Morton 1984; Forde 1986), Many
of the shrubs (e.g. RAagodlia spinescens, Enchylaena
(tomentosa and Chenopodium gaudichaudianurn)
are found at higher frequencies underneath trees
and large shrubs than in the open (c.g. Barker
19723), Two hypotheses have been proposed to
account for this. First, the clumped distribution
reflects the pattern of dissemination by birds, the
hirds defaecating seeds while perched in trees (e.g.
Osborn et al. 1935; Leigh & Mulham 1965; Forde
1986). Alternatively, or in addition, Barkei (1973)?
suggested that the clumping was due to mote
favourable growing conditions beneath {ree
canopies, To distinguish between these rwo
hypotheses requires measuring the seed rain bencath
* Department of Botany, University of Adelaide, ¢9.P43.
Box 498, Adelaide, S. Aust. 5001.
Department of Zoology, University of Adelaide, O.P:0.
Box 498, Adelaide, 8, Aust, 5001,
~ Present address; Botany School, Downing Street,
Cambridge, CB2 JEA, England,
Present address: Facultad de Silviculiura y Manejo de
Recursos Renovables, Universidad Autonoma de Nuevo
Leon, Apdo. Postal 104, 67700 Linares V.L,, Mexico.
Barker, 8. (1972) “Effects of Sheep Stacking on the
Population Structure bf Arid Shrublands in South
Abstralia™, Ph.D. thesis. University of Adelaide,
Adelaide, Unpubl
wd
and between Gee canopies, and determining rares
of germination and seedling establishment in the
different microenvironments.
In this paper, we (1) document the distribution
of several species of shrubs in relation to: tree
canopies on Middleback Station, 27 km north-west
ol’ Whyalla; and (2) report the use of a “seed trap”
to measure the sedd rain. beneath trees due to birds
and discuss the data so derived.
Materials and Methods
Study site
The study site was in the south-western corner
of Overland Paddock, Middleback Sration
(32°57'S, 137°24'E) in an arca of open woodland
which is moderately grazed by sheep. Two tree
species predominated: Myoporum platycarpun
R.Br. which grows to 10 m and has a thin elevated
canopy: and Acaciu pupyrocarpa Benth, which bas
a dense low spreading canopy. The two Irees. were
infected by mistletoes: Af, plaiycarpum by Lysiene
exocurpi (Behr) Tiegh, and A. papyrocarpa by
Amyena quandang (Lindl). Teigh. The understorey
was dominated by the chenopods, Maireana
sedifolia (F Muell.) P. G. Wilson, Atriplex vesicaria
Heward ex Benth., Malreana prramidata (Benth.)
P. G. Wilson and Rhagodia ulicina (Gand,) P. G.
Wilson, Four species of small shrubs (Rhageadia
Spinescens var, spinescens F.Muell., Chenopodium
gaudichuudienum (Mog.) BP. G. Wilson, Enevimerne
tomentosa R,Br., Lycium australe F.Muell.), the nwa
2 M_ TESTER £&T AL.
Tistletaes and two small trees of large shrubs,
Exocarpus aphyllus R.Br, and Meterodendrum
vleaejolium Desf, produced fruits that were
consumed by birds in the area (Reid 19844; Forde
LYR6).
Distribution of stitubs
Distribution of shrubs in three zones; under the
canopies of M, platycarpum, under the canopies
of A. papyrecerpa, and in the spaces between the
trees and large shrubs was measured, Interspaces
were at least 2m from the canopy of any tree or
large Shrub. Two hundred 0.5 m0.5 m quadrars
were placed haphazardly in cach zone, and the
presence of a shrub species in each quadrat was
seored if any part of its canopy occurred in the
quadrat, The diameters of the canopies of 30
individuals of each shrub spevics in each zone was
Measured by measuring the width across the plant
along two axes (north-south and cust-west). Since
the mean canopy diameter of a particular shrub
Species was the same under the two tree Species as
well as in interspaces, Incidence in quadrats provides
2 measuee of the relative density of a plant species
in €ach of the three zones. However, the canopies
of ihe different shrub species were not the same
diameter (Table 1), so the relative densities of the
different species cannot be deterniined from the
quadrat frequencies,
+ Reid, N. (1984) “The Role of Birds in the Reproduction
of an Arid Zone Popiilation ofGrey Mistletae 4 myena
quandang (Loranthaceae),” Ph.D. thesis. University of
Adelaide, Adelaide. Unpubt,
Trapping for bird droppings
Twenty-two traps to collect bird droppings were
deployed in March 1984. Traps were made by tying
1.2145 m sheets of terylene voile (mesh 0.8x
().3 mm) Lo steel stakes to form a catching area of
1.0%1.5 m that stood 90 cm above the ground (e.g.
see Sorensen 1981). This was the largest size of trap
that could be conveniently erected under the
canopies of the trees. A stone was placéd ti the
centre of the catching arca tu prevent spillage of
seeds by strong winds, liqual numbers of traps were
Placed beneath canopies of mature M. platycarpunt
and mature 4. papyrocarpa. All of these trees were
infected with varjable amounts of mistletoes. Traps
were inspected and cmptied of seeds at Iwo
4-monthly intervals. The seeds were identified using
a4 reference collection of seeds compiled in the area.
Many of the traps were damaged 4 to § months after
deployment and observations ceased after 8 months.
Results
Distribution of shrubyspecies in Overland Paddock
Nine species of shrubs were detected in the six
hundred 0.5.0.5 m quadrats. OF these, five species
occurred significantly more frequently under ihe
canopies of either Acacia papyrocarpa ar Myo-
porum platyearpum than in the interspaces between
the trees and large shrubs (Table 1). Of these, three
species produced fleshy fruits that were dispersed
by birds (Chenopodium gaudichaudianum,
Enchylaena tomentosa and Rhagodia spinescens),
while the other two species (C. desertorum and
Tanve L. Frequency of accurrence of shrubs in 200 eae (0.5 *0.5 m) in each of three zones in Overlunad Paddoce,
Mareh 1984.
—_ i
Frequency in 200 quadrats
Meun canopy" heneath beneath in x
Shrub species area (m>) A, pupyrocarpa = M. plalycarpnim interspace valine
Atriplex vesicaria 40 38 74 44 14.3 *4*
Chenopadium desertor yim os 12 10 2 6.0 *
C. paudiéhaudianum OAT Sl 15 () 60.9
Enehylagna tomentosa’ ad 32 34 7 18. ***
Lycium australe’ 49 7 a 13 2,2 NS
Maireana pyranudata — i i 1] Q.0 NS
M. sedifolte 42 42 4 LIONS
Rhegodia spinescens* — 4] 22 0 40.) 44
R. ulicinea — 23 2 18 4.1 NS
A ee
* Canopy areas calculated from diameters of 60 or 90 individuals dependiag on sumber of zones in which plant species
occurred, Standard errors were Jess than 30% of the mean and there were no significant differences in the canupy
area OF it shcub species in different zones (ANOVAs, P=0,05), so dali for the different zones were pooled,
he oo tia (2 ef) tests the null hypothesis that shrub species were evenly distributed across Ure three zones:
"(1,05
F* P<
ete P(N
NS not significant
“Shrubs that produce flashy fraits consumed by birds,
SEED DISPERSAL BY BIRDS AND PLANT DENSITY 1
Airiplex vesicaria) had wind or ant-dispersed seeds.
Enchylaena tomentosa and C. deserrorum were
equally frequent under both canopies, but 4,
yesicarid was encountered most frequently under
Mf. platycarpum, and C, gaudichaudianuen and R.
spinescens were encountered most frequently under
Acacia papyrocarpa.
Four species of shrub showed no significant
clumping under the canopies cf trees (x7 tests,
p>0.05). Of these, the Malreana species and R.
witcina have wind or ant-dispersed seeds but Lrefuen
rustrale produces fleshy fruits that are dispersed by
birds and perhaps reptiles.
Seed rain detected by traps
Table 2 lists the quantity of seeds collected in the
traps'set under the canopies of Acacta papyrocurpa
and M. platycarpum during two time periods, The
numbers of seeds deposited per tap was highly
varlable, with variances often exceeding the means,
There was no significant difference in the tatal
numbers of sceds deposited under A. papyrocarpa
and M, platycarpum (rank-sum tests, p>0,05),
More seeds were caught during the period July-
November than between March-July (rank-sum test,
p<0,002), In general, the number and species of
seeds being deposited during each period reflected
the fruiting seasons of the plants, Chenopodium
gavdichaudianum, Enchylaena tomentosa and
Exocarpus aphyllus tryited mainly during winter
and early spring, while Heterodendrum olesefoline
and Lyciven australe fruited mainly in aeumn,
Rhegodia spinescens fruited mainly during summer,
but like Enchylaena tormentosa and Exocarpus
aphyllus, produced small quantities of fruit
thronghout most of the year:
Many of the traps also collected seeds of misthe
toes, These data are not presented since mistletoe
seeds must be deposited on the branches of host
(recs fo be effectively disseminated, The seeds
collected in the traps therefore represented
ineffective dispersal.
Discussion
The purpose of this study was to measure the
distribution of shrubs, particularly the fleshy-fruited
chenopods, in telation to tree cover, and to
delermine the usefulness of traps in measuring seed
rains, Prior to this study, staremenrs. that fleshy-
fruited shrubs were clumped under trees were based
largely on qualitative observations (e.g, Osborn et
al. 1935; Leigh & Mulhaim 1965; Forde 1986). Our
quadrat data show that three feshy-fruited shrubs
were found mainly under trees, and that the extent
of this clumping varied from species to species.
Rhagodia spinescens and Chenopodium
gaudichaudianum were almost exclusively found
under tree canopies, but more frequently under
Acacia papyrocarpa than under Myoporurn
Platvearpum, Enchyluena tomentosa was foulid
with equal frequency under both tree species and
occasionally in interspaces, while Lyeium australe
was cqually abundant under trees and in interspaces
(Table. 1). However, clumping beneath trees. was n0t
restricted (© bird-dispersed ‘species. Among the
shrubs dispersed by wind or ants, A/riplex vesicoria
was more Common beneath M. plalycarpurn than
Acacia papyrocarpa or in interspaces, and C.
deseriorum was significantly more common under
both trees than in the interspaces. Higher densities
of shrubs under trees could occur because (1) more
seeds are dispersed beneath trees and/or (2)
germination rates and seedling establishment are
Tani e 2, Nuenbers of seeds deposited by birds in traps set under Myoporum platycarpum and wider Acacia papyro-
carpa in Overland Paddock under two time periods in !984-
Species of shrub
raps set under M, platvcarpuni
Chenopodium gaudichaudianum
Envhyluena lomentose
Exacarpos aphyltus
Heterodendrum oleaefolium
“Lbevaen australe
Rhugodia spinescens
Total
Traps set under 4, papyrocurpu
Chenopodium goudichaudianum
Eachyluena tomentosa
Exocarpus aphyllus
Mererodendrum oledefalijint
Lycium australe
Rhagudta spigesceny
Total
Mean number of séeds per trap+S.E-
March-Jily Jyly-November
n=11 t+
0.5 40.2 48+ 33
2.34 2.0 (O84 3.5
3.7 +31 23.5+10.9
3.271.4 ad: a0
D.4+0,2 101 0.0
1bL0,7 02+ OO
12.99 7,2 39,04 7.)
i-4l n=8
&.040.0 a+ 04
14+60.7 §R+ 97
5.84138 2aAt 92
L.740.8 06+ 0.5
0,2=02 O04 0,0
0.4404 blz i
9.5444 36.2+14.2
4 M_ TESTER £7 AL.
higher under (revs than in the open (c.g. Barker
19723) Forde 1986). These Iwo eaplanations are
sequential and wot mutually exclusive, Both are
prubably important in determining the distribudions
of various fleshy-fruited planre in Overland
Paddock. The first thing ro demonstrate, however,
is Whether birds disperse seeds mainly to sites under
Irets.
On Middieback Station, Meshy-lruited shrubs
have their seeds dispersed largely hy birds anil most
bf the seeds appear to land under trees. Although
our [raps Were not deployed ta denionsirate thal
more seeds were deposiced underteees than in the
open, other observations enable such a conclusion.
Ju Overland Paddock, two species.al honeveater
are largely responsible for the dispersal of seeds by
bitds, the Spiny-cheeked (Acunthugenys rufo-
gutoris) and Singing (/ichenoastamns virescens)
Honeycaters (Reid unpubl.). These honeyeaters
oiainly feed on the nevtar and fruit of mistletoes,
supplementing these foods with insects and fruits
of other shrubs (Reid 1984}. More than 90% ol
their facces are discharged while the birds are
perched more than a metre above the ground in
trees or tall shrubs (Reid unpubl.). Consequently
bird-disseminaled seeds should be concentrated
under the canopies of trees.
‘The traps that we deployed under the trees were
successful in collecting seeds, and the quantities
collected {10-30 seeds/m*/4 months; Table 2) are
consasient with eslimates of the mimbers of seeds
(hal should have been voided by birds. Assuming
that chere were 1.2 hotecyeaters/ha, that these
joneyeaters consumed 8O-L6C seeds from fleshy-
fruited shrubs per day, that the birds largely
disseminatet these seeds under (he canopies of trees
und that the tree cover was 10%, then in 4 manths
approximately 10-20 sceds/m* should have been
voided by birds under trees (Reid 1984", in prep.),
The numbers of seeds collected by individual
taps were highly variable. ‘Two potential sources
of variation probably account for this; variation
in the ise oF trees or parts of trees by birds, and
variation in faceal composition, Certain irees or
paris ol trees may be used more extensively by hirds
because of their location, architecture or the
amount of food associated with them, Trees
centrally-locwted in a bird's territory or near the
bird's nest may be used more [requently (e.g.
Qrians & Pearson 1979), The densily of the foliage
or height of a lree may provide birds with better
pPrifection Fram exposure or predators, or better
varrtage points for detecting predators or intruders,
and so trees with these attributes may be used mare
Irequemily, The trees in the study area also differed
1 The NUMbers af Mistleines (hey supported (Rei
19849) and probably also in the quantities of fleshy
(fruits provided by the shrubs heneath them, Birds
should spend more time in trees where the food
supply is greatest (e.g. Charnoy L976; Pyke e/ af.
L977). The second major source of variatien
concerns the fagces of the birds, A single faecal
dropping may contain 0 to 20 seeds, and much of
the variation belween traps may be due to varia-
tions in the numbers of seeds in a faecal dropping
rather than variations in the use of trees by birds.
We counted the total number of seeds in the traps,
not the number of Facces. Future studies may wish
to control for these variations, or at least accent
for them by recording the architectural features,
the locations, arid associated food supplies of the
trees under which the traps were placed, Use of
several traps under the same tree also would allow
within-tree Variation in bird use to be measured,
The size and design of the traps seems appro-
priate for measuring seed rains under trees in the
arid zone. Traps, however, would need to be set
closer to Lhe ground and under shrubs if seed rains
were to be measured in the interspaces becween
trees, because the honcycaters only defaecate while
perched (Reid 19844), The size of the traps
(1.5 x 1.0 m) also seems appropriate since larger
traps would have been tuo awkward to erect under
trees and the size was such that there was a
reasonable chance thal a trap would collect at least
some seeds, Traps failed to collect sceds on only
SIX Occasions out of 34 four-month exposures.
Based on our experience, we wauld recommend
clearing traps of their seeds every two months, and
using @ stronger mesh for ihe catching material.
This would reduce the loss of farcul material duc
to the cutching material deteriovating with (inie. We
found that terylene voile deteriorated after about
four months, particularly under Mlyoporum
plarvearpum where Lhe raps were exposed tn mare
sunlip tit
Given (hal most of the seeds are dispersed to sites
uncer trees we would expect bird-dispersed plants
to be most abundant under trees in the arid vone.
[it gesteral this is die case, bul lhere ate exceptions.
Lyeiunt ausirafe was equally abundant under trees
and in interspaces, and the low clonal tree, Hevero-
dendrum olewefuliam, often occurs away from
other trees. These species reproduce extensively by
rool suckers and are apparently Iong-lived relative
io the trees (Purdie 19695; DB. E, Symon pers.
comm.). Lith: is known about their seedling
recrulfment (cf. suckering), since seedlings are
tarely Seen: no seedlings of A; vleaefolium have
3 Purdie, R. (1969) “The Population Steucture of
Seleered Arid Zone Tree Species”. B.S. (Mutts) (hesis.
Univ, of Adelaide. Unpubl.
SEED DISPERSAL DY BIRDS AND PLANT DENSITY $
been detected on Middleback Station over the last
20 years (Lange unpubl.). Seedlings of these spocies
may only establish under trees, but once esta-
blished, may produce suckers to exploit interspaces
between trees where competitive interactions with
the established trees presumably are reduced, The
present distribulicn of these plants, then, may oot
reflect the sites where scedlings established, and the
relatively high densities of these plants away from
trees is not. necessarily inconsistent with most of
the seeds being deposited under trees. In addition,
Lyeivm australe produces fruit close to the ground
and lizards may therefore consume and disperse
some Lyeium secds to interspaces.
The best evidence thal factors other than seed
dissemination by birds are also involved in the
eventual pattern of distribuuon in these Fleshy-
ruited plants comes from comparing the distri-
butions of the three berry chenopods, Rhagedia
spinescens, Chenopodium eaudichaudianum and
Enchylaena tomenresa. These three species are
almost exclusively bird-dispersed, and similar
quantities of seeds were deposited under 4. papyro-
carpe and M. platyearpunt (Table 2), The density
of Enchylaena tomentosa under thetwo tree species
was similar, and so the pattern of seed dispersal
by birds may provide an adequate explanation for
its distribution. However, both RX. spinescens and
C. gaudichaudianur had higher densities under A.
papyrocarpa than M, platyearpum. Factors besides
dispersal by birds are needed to explain ukis. The
most likely explanations involve the climatic or
edaphic differences encountered beneath trees,
coupled with differences in the regeneration niche
of the shrubs. Acecia papyrocarpa and M_ platy-
cerpunt differ in architecture (branching patiern
ef.) and hence in the amount of shade they provide
a germinating seed or scedling. The canopy of A.
pepyrocarpu is relatively dense and provides
shade and w thick cover of litter. Myoporam
Platyearpum has a tigh thin canopy, so plants
beneath il are subject to conditions of more severe
water stress (Barker 1972+). Other factors such as
increased ‘nutrient cycling under A. papyrocarpa
(Correll 19675} of increased grazing pressure under
M. platycarpum could also be involved. Similar
sorts of factors wauld account for the fact. that
Afriplex vesicaria and possibly Rhagodia ulieina
were more abundant under Mf. platyearpum than
A, pap procarpa, Careful field experiments will be
required to determine the factors (edaphic, climatic,
biotic) that are important for the germination and
seedling establishment of each species. These
experiments should involve transplanting seeds or
seedlings between the canopies of differcnt tree
species as well as info the interspaces. Only after
these experiments have been performed will we
have Ihe necessary evidertee to state the relative
vontributions of bird dissemination of seeds and
environmental factors in devermining the distri-
bution of Fleshy-Pruited shrubs in the arid zone.
Acknowledgments
We would like to thank Tim Croft and Christina
Morris for assistance jn the lleld, and Andrew,
Lesley, Don and Penan Nicolson for their generous
hospitality at Middleback Station, Hugh Ford and
an anonynious reviewer provided helpful comments
on an earlier draft, This work was based at the
Middleback Field Centre which is supported by
Broken Hill Propriety Co, and Mitsubishi Motors
(Australia) Ltd.
© Correll, R. L. (1967) “Studies on the Nitrogen
Economy of Semi-Arid Vegetation at Yudnapiona
Station, South Australia.” M.Se, thesis. University af
Adelaide, Unpubl,
References
Cratwoyv, E. L. (1976) Optimal foraging: de marginal
value theotem. Theor, Pop, Biol, 9, 129-446.
Davinson, O. W., & Morton, S_. RK. (1984) Dispersal
adaplations of some Acecia-ypecies in Une Australian
urid zone. Lenlogy 65, 1038-1051.
Forpr, N. £1986) Relationships between birds and trnits
in tomperate Australia, pp, 42-58, fr 1. A. Ford &
D.C, Paton (Eds) “The Dynamic Partnership: Birds
and Plants in Southern Australia.” (Govt Printer,
Adelaide),
Hakeenk, J. L. 11977) “Population Biology of Plants.”
(Academie Press, London),
Letch, J. A, & MULHAM, W. E. (1965) “Pastoral Plants
of the Riverine Plain,” (Jacaranda Press, Melbourne).
Ornians, G. FL, & Peansas, N. E. (1979) On the
theory of cencral place foraging. pp. 155-177, In D.
J, Horn, R, Mitchell & G, R, Stair (Eds) “Analysis of
Ecological Systents." (Ohio State Univ. Press,
Columbus, Ohio).
Osporn, T. G. B., Woon, J, G., & PALTRIDGE, T. B.
(1935) On the climate and vegetation of the Koonamore
Vegetation Reserve ta 1931. Prac. Linn, Sac. NWSW.
Gl), 992-427,
Pyat, G. H., Puroiam, H-R., & Cnannoy, EL. (1977)
Optinial foraging @ selective review Of theory and lests,
O. Rev. Bini. 32, 137-154.
SORENSEN, A. EB. (1981) Interactions between birds and
fruit in a temperate woodland. Oecofogia 50, 242-249,
A PROPOSED REFERENCE SECTION FOR THE TORTACHILLA
LIMESTONE
BY A. S. BEECROST*, J. H. CANN? & C. STOCKSIEKE
Summary
Roadside outcrops of sedimentary rocks near Port Noarlunga, South Australia, are identified as Late
Eocene stratigraphic units of the Noarlunga Embayment within St Vincent Basin. Fossil planktonic
and benthic foraminifera from these rocks compare favourably with stratotype Tortachilla
Limestone and the Tuketja Member of Blanche Point Formation. An exposure of highly
fossiliferous, glauconitic calcarenite, bounded below by the South Maslin Sand Member of Maslin
Sands, and above by the Tuketja Member of Blanche Point Formation, is proposed as a reference
section for Tortachilla Limestone.
KEY WORDS: Eocene, foraminifera, Noarlunga Embayment, St Vincent Basin, Tortachilla
Limestone.
A PROPOSED REFERENCE SECTION FOR THE TORTACHILLA LIMESTONE
by A. S. Bernesoery J. AH. CANN] & C, STOCKSIEKT
Summary
Beeckarr, 4.5, Cann, dH. & Srocksirk, © (1987) A proposed reference section for the Torlachitta
Limestone, Jrans, R, Sac, 5. Aust, W1(1), 7-15, 29 May, 1987,
Rowdsidle outcrops of sedinientary tocks near Port Noarlunga, South Austratia, are identifiod as Late
Cocene stratigraphic units of the Noarlunga Embuyrnent within St Vincent Basin. Fossil planktonic and
bentine foraminiena fron these racks compare favourahly with siratorype Tortachilla Limestone anit (he
Tuketia Member of Blanche Poin! Formation, Anesposure of hizhly lossiliferous, glauconitic calearenite,
bounded below by the South Maslin Sand Member of Maslin Sands, and above by the luketsa Member
of Blanche Poin Formation, is propased as a veterence section for Tortachilla Limestone,
Key Worbs: Eocene. foraminifera, Noarlunga Embayrment, St Viruent Basin, Torrachitla | amestone.
Introduction
The ‘fortachilla Limestone is a Late Eocene
stratigraphic Member of St Vincent Basi (Reynolds
1953; Stuart 1970; Buonaiuto 1977; Cooper 1979;
Beecroft 1980): Jenkins er al 1982). Within the
Willunga Embayment, at the type locality, Maslin
Bay, the Tortachilla Limestone bas a maximum
thickness of about two metres. Mt overlies the poorly
fassiliferous. South Maslin Sand Member of the
Maslin Sands (Cooper 1979) and is in turn overlain
by (he younger Tuketja Member of the Blanche
Point Formation (Jenkins ef.a/. 1982), Buonaiuto
(1977) confined the Tortachilla Limestone to the
lower Polyzoal Limestone Member of Reynolds
(1953) and this was follawed by Cooper (1979),
though not by Jenkins er i (1982), whase
nomenclature is used in this paper. Fig. 1
summarizes this stratigraphy,
About (en years ago the beach at Muslin Bay was
proclaimed available for nude bathing and this
factor has deterred some study groups [rom visiting
the stratotype area. Also, an equivalent section at
Whitton. Bluff, Christies Beach, is no longer
accessible for study due to coastal protection works.
There is therefore a need for nominution of other
ourcrops as reference sections of the Tertiary units.
* Peparrmeni of Geology, University of Adelaide, G,P.O,
Box 498, Advlaide, 5, Ausr. 5001.
PAshool of Pure und Applied Sciences, SALCLALE.
(Salisbury Campus), Smith Road, Salisbury Fast,
5. Aust. S109,
+Cenve lor Environmental Studies, Universin of
Adelaide. G.P.O. Box 498, Adelaide. 5. Ausr, SOO,
' Beewoltt, A. S. (1980) boraminiferal bioficies of the
Tortachilla Limestoue and the Blanche Point bormation,
Late Locene, Willunga Sub-basin, South Australia,
HOA.(Hoas.) thesis, Univ. of Adeliido,
Outcrop descriptions
Daily (1952)? soted outcrops of Tertiary
sediments of the Noarlunga Embaymient in ctiffs
and road cuttings adjacent te the lower
Onkaparinga River. More recently, additional road
works have exposed these strata in greater detail
(Stocksiek 19834). Aq exposure on River Road
(Fig. 2, Site 1) presents more Can HX) metres of
easily accessible and continuaus outcrop. Part of
this exposure is shown in Fig, 3.
From road level, ycllow. brown clays give way to
coarse, limonitic, crossbudded sandstone, apparently
non-fossiliferous, These sediments have a thickness
of about 2.5 metres and are characteristic of the
South Mastin Sand Member, Overlying, Tortachilla
Limestone, typically glauconitic green, sandy arid
richly fossiliferous, is ) 2 metres thick. Glauconitic
tharls of the Tuketia Member form the upperinest
beds of the outcrop. These three outcropping units
are essentially similar in lithological detail (o
descriptions of stratutype material given in Cooper
(1979), Beeguse of ease of access and clarity of
stratigraphic boundaries, this exposure 14 here
proposed as a reference section for the Tortavhilla
Limestone. Fossil foraminifera trom the section are
discussed later below,
In roadside ooterop west of the Reference Sectivn
(Fig, 2, Site 2) only 0.5 metreol South Maslin Sand
Member appears wbove toad level. Tortachilla
Limestone is bleached and its upper boundary is
indistinct. Tuketja Memiber constitutes most of the
outcrop, though the higher, harder, more prominent
beds may represent the Gull Kock Member of the
* Daily, H. (1952) Stratigraphy and geology of the
Noarlunga basin. B-Su(Hons.) thesis, Univ. of Adelaide,
+ Stocksiek, C. (1963) Some observations of (he Tertiary
stritta Oulcropping adjacent to the Onkaparinga estuary,
South Australia, Geology project report, 8. Aust. Coll.
Adv Edue, Salisbury
§ A. S. BEECROFT, J. H. CANN & C, STOCKSIEK
SUMMARY OF TERTIARY STRATIGRAPHIC NOMENCLATURE
TATE 8
BURR | oENNaNT | REYNOLDS | CRESPIN | COOPER [JENKINS etal
1846 1896 1953 1954 1979
PORT ALDINGA
WILLUNGA (
LIMESTONE
BEDS
AvoINGa | >
MEMBER
CHINAMAN GULLY BEGS CHINAMAN GULLY FOR! INAMAN GULLY FORM
SOFT MARLS
PERKANA
MEMBER MEMBER
GULL
ROCK
MEM&ER
PORT WILLUNGA FORMATION
PIRRAMIMMA SAND MEMBER
OLIGOCENE
BLANCHE
BANDED POINT
MARLS LIMESTONE
MEMBER
(NOARLUNGA
TRANSITIONAL LIMESTONE)
MARLS MEMBER
GLAUCONITIC
LIMESTONE
MEMBER
GULL
ROCK
MEMBER
BLANCHE POINT FORMATION
TUKETJA MEMBER
BLANCHE POINT FORMATION
LATE EOCENE
TORTACHILLA
oD
®
c
am
wm
w
o
w
w
™&
3
co)
2
=
Se
wo
a]
ce)
Cc
POLYZOAL TORTACHILLA
LIMESTONE LIMESTONE | LIMESTONE
MEMBER
EOCENE
TORTACHILLA LIMESTONE | BLANCHE POINT MARLS
TERTIARY
SOUTH SOUTH
SOUTH MASLIN MASLIN
MASLIN SAND SAND
SANDS MEMBER MEMBER
MASLIN
SANDSTONE
NORTH
NORTH MASLIN
MASLIN SAND
SANDS MEMBER
MIDDLE EOCENE
Fig. 1. Summary of stratigraphic nomenclature for St Vincent Basin, South Australia. No scale implied.
REFERENCE SECTION TORTACHILLA LIMESTONE 9
Oval
PORT
NOARLUNGA
~
i
D
S
s
N
4
S
S
O 500
pe
METRES
COMMERCIAL
CHRISTIE
ADELAIDE
I
— 1
Gulf St Vincent
Fig. 2. Location map showing sites of outcrop of Tertiary sediments referred to in text.
the Blanche Point Formation. The sequence
apparently dips gently westwards.
At the intersection of Morton and River Roads
(Fig. 2, Site 3) Tortachilla Limestone outcrops
prominently. It is conspicuously fossiliferous (Fig. 4)
and overlain by both Tuketja and Gull Rock
Members of the Blanche Point Formation.
Foraminifera
Microscopic examination of washed material
from the identified Tortachilla Limestone at Site 1
revealed a rich micro-fauna dominated by benthic
foraminifera, with occasional ostracods. Scanning
electron photomicrographs of some species of the
foraminifera are shown in Fig. 5. A similar
OS
10 A. 8. BEECROFT, J. H, CANN & C
-, STOCKSIEK
* a
Nig. 3, Composite photography of part of the proposed reference section for the Tortachilla Limestone.
foraminiferal assemblage occurs at Maslin Bay in
stratotype Tortachilla Limestone and has been
referred to Zone PIS (Lindsay 1981.4 and refs,
therein),
Significant planktonic foraminifera present
include; Subbotina linaperta, 8. angiporoides,
Tenuitella aculeata, T. gemma, T. inselita,
Turborotalia nana, T, centralis, T) cerrouzulensis,
Chiloguembelina cubensis, Pseudohastigerina micra
and Globigerinatheka index. The lower disjunct top
lo Tenuitella aculeata occurs within this unit, while
Turboratalia cerroazulensis has only recently been
found in the type section (Lindsay 1981) and is not
known With any certainty to occur locally above the
fortachilla Limestone.
Among the benthic foraminifera, the presence of
Pseudopolymorphina carteri, Linderina glaessneri
and Maslinella chapmani is significant.
Pseudopolymorphina carteri is restricted in
occurrence tO the Tortachilla Limestone and basal
Blanche Point Formation (Tuketja Member) at
Maslin Bay, while the presence of Linelerina
&laessneri at this level represents an extra-tropical
excursion (Lindsay 1967, 1969; McGowran 1978),
The presence of Muslinella chapmani appears to be
lemperature controlled, being found in the
Tortachilla Limestone and Tuketja Member, and
then not again until the basal part of the Aldinga
Meinber of the Port Willunga Formation (P14)
where L. g/aessneri also reappears bnetly,
‘Lindsay, J. M. (1981) tertiary stratigraphy and
foraminifera, Adelaide, South Australia. M.Sc. thesis,
Univ. of Adelaide.
The remainder of the assemblage are specics
which cover a wider stratigraphic range, but as an
assemblage, the unit contains both planktonic and
benthic elements which are consistent with
stratotype Tortachilla Limestone.
The overlying unit contains rare, small, broken,
but nonetheless unambiguous Hanikenina
primitiva, which unequivocally confirms that this
unil is the Tuketja Member of the Blanche Point
Formation, and hence supports the identification
of the Tortachilla Limestone unit. The Tuketja
Member here does not contain 7! weuleata nor L.
glaessneri, consistent with the type section at
Maslin Bay, although P carleri and M. chapmani
are present in reduced numbers.
Conclusions
The presence of key species, in particular
Tenuitella aculeata, Turborotalia cerroazulensis, T.
centralis, Pseudopolymorphina carteri, Maslinella
chapmuni and Linderina glaessneri enable this unit
to be recognized stratigraphically as Tortachilla
Limestone, Identification is further emphasised by
the presence of Hantkenina primitiva in the
overlying unit, which is thus confirmed as being the
Tuketja Member of the Blanche Point Formation,
The foraminiferal fauna of the Tortachilla
Limestone enables correlation with the equivalent
level in the Browns Creek section of south-western
Victoria and the Nanarup Limestone of the Bremer
Basin, Western Australia (McGowran & Beecroft in
prep.),
REFERENCE SECTION TORTACHILLA LIMESTONE Il
my A
Fig. 4, Selected fossils, photographed at outcrop of the Tortachilla Limestone at site 3. A and D, bivalves; B, brachiopod;
C, gastropod; E and F, bryozoa. all x 1.5 approx. (diameter of Aust. 20 cents coin is 28 mm).
Me
I
n
4
U
F
6)
os]
Zz
Z
<
O
x
=
=
the
oO
w
OU
ira
a)
wi
<
REFERENCE SECTION TORTACHILLA LIMESTONE 13
Acknowledgments
Sharon Proferes drafted figures 1 and 2; Richard
Barrett provided photographic assistance; Brent
Bowman picked the Hantkenina primitiva; Chris
Moore typed the final manuscript. We thank Dr
Richard Jenkins and Mr Murray Lindsay for
critically reading the manuscript.
References
BuonaluTo, M. F. (1977) Revision of the Australian
‘Tertiary species ascribed to Limatula (Wood) (Mollusca,
Bivalvia). Trans. R. Soc. S. Aust. 11, 21-33.
Coorer, B. J. (1979) Eocene to Miocene stratigraphy of
the Willunga Embayment. Rept. of Investigations 50,
Geol, Surv. S. Aust.
JENKINS, J. B., McGowran, B., BEECROFT, A. 5, &
FitzceraLp, M. J. (1982) Lithostratigraphic
subdivision of the Blanche Point Formation, Late
Eocene, Willunga Sub-basin. Quart. Geol, Notes, Geol.
Surv. S. Aust. 84, 2-7.
Linosay, J. M. (1967) Foraminifera and stratigraphy of
the type section of Port Willunga Beds, Aldinga Bay,
South Australia. Trans. R. Soc, S. Aust. 91, 93-109.
__ (1969) Cainozaic foraminifera and stratigraphy of
the Adelaide Plains Sub-basin, South Australia. Bull.
Geol. Surv. S. Aust. 42.
McGowran, B. (1978) Early Tertiary foraminiferal
biostratigraphy in Southern Australia: a progress report.
BMR Bull. 192, 83-95,
ReyNotps, M. A. (1953) The Cainozoic succession of
Maslin and Aldinga Bays, South Australia. Trans. R.
Soc. S. Aust. 76, 114-140,
Stuart, W. J. (1970) The Cainozoic stratigraphy of the
eastern coastal area of Yorke Peninsula, South Australia.
Ibid. 94, 151-178.
Fig. 5. Selected foraminifera from the Tortachilla 1.imestone at the proposed reference section. A, B Turborotulia
verroazulensis (Cole) x 75; C, D Tenuitella aculeata (Jenkins) x 220; E, F Hantkenina primitiva Cushman and
Jarvis x 120; G Globigerinatheka index (Finlay) x 145; H Subbotina angiporoides (Hornibrook) x 220; 1, J
Chiloguemtbelina cubensis (Palmer) x 220; K, L, M Pseudohastigerina micra (Cole) « 145, N, O T. insolita Jenkins)
x 220, P Pseudopolymorphina carteri Quilty x 27; Q Stomatorbina concentrica (Cushman and Bermudez)
x 65; R, S Eponides repandus Fichell and Moll) x 45; T Linderina glaessneri Quilty x 27.
A COMPARATIVE ANALYSIS OF EOCENE/OLIGOCENE BOUNDARY
OSTRACODA FROM SOUTHEASTERN AUSTRALIA AND INDIA
WITH RESPECT TO THEIR USEFULNESS AS INDICATORS
OF PETROLEUM POTENTIAL
BY K. G. MCKENZIE* $ D. K. GUHAT
Summary
Eocene/Oligocene boundary Ostracoda were analysed from selected wells in the Adelaide Plains
Sub-Basin, South Australia and Cambay Basin, India. Source-rock characteristics of the sequences
were determined - based mainly on ostracode parameters, with some additional information coming
from their glauconite and gypsum content. Numerically similar ostracode counts were made for
both sets of samples. The parameters studied were: carapace/valves ratio; adults/juveniles ratio;
percentage of fragments; percentage of crushed and worn specimens; Krithe type; percentage of
pyritised specimens.
Results (which concur with exploration results to date) indicate that the Eocene/Oligocene boundary
zone sediments have little petroleum potential in South Australia, but high potential in India. This
conclusion was largely reinforced when the South Australian borehole Ostracoda were analysed in
more detail. Consistent results were also obtained when the same parameters were determined for
Ostracoda in outcrop samples collected from Aldinga Bay, South Australia.
KEY WORDS: Ostracoda, petroleum indices, Eocene/Oligocene boundary, South Australia, India.
A COMPARATIVE ANALYSIS OF EOQCENE/OLIGOCENE BOUNDARY OSTRACODA
FROM SOUTHEASTERN AUSTRALIA AND INDIA WITH RESPECT 10 THEIK
USEFULNESS AS INDICATORS OF PETROLEUM POTENTIAL
by K. G. McKen/ie*t & D, K, GuHAt
Summary
MeRenzin, Keo & Gata, De oR, (L987) AN vonipacdhive analysis Of Locene/Oligocene boundary
Osrracoda from southeastern Australis aod India with respect to their usefulness as indicaters of
petrotcum porenual, Tray. R. Soe. S. Aus 1110), 15-23, 29 May, 1987,
Eocene/Oligocene boundary Ostracoda were analysed from selected wells in the Adelaide Plains Sub-
Basin, South Australia and Cambay Basin, India. Source-rock characteristics of the sequenves were
determined — based mainly on ostracdde parameters, with some aitditional mformation coming from
their glauconite and gypsun) content, Numerically similar oxtracode counts were made for both sets of
samples. The parameters studied were: curapace/ valves ratio; adults/(uveniles ratios percentage of fragments:
percentage of crished and work specunjens; Arithe type; percentaye of pyrizised specimens.
Results (which concur with exploration results to date) indicate thar the Baceney Oligocene boundary
vong sediments have litle petroleum potential in South Australia, but high potential in India. Vhrs canelusian
was largely reinforced when the Saurh Australian borehole Ostracoda were analysed in mare detail, Consistent
fesults were also ohianed when the same parameters were determined for Ostracoda in outcrop samples
collected from Aldinga Bay, South Ausoratia.
Ky Words: Ostracoda, petrolcun indices, Eovene/Oligocene boundary, South Australia, India.
Introduction examination ol numerous
During September-October 1983, one of us
(U.K.45.) visited Austraha under the aegis of the
Australia-India Science and Technalocy Agreement
to study the Tertiary ostracode microfaunas of
southeastern Australia for comparison with
ostracodes in Indian Tertiary segucnees. Lhe senior
suihor (&.G.M.) acred as host for the visit und a
ty-operative project was. initiated,
We soon decided on the EocenesOligocene
boundary zone because it was well understood bath
in India and Australia and was known to be
important for petroleum exploration in many parts
of the world (Potmerol & Premoli-Silva 1986),
including India (Guha & Pandey L980), Australia
(Douglas & Merguson 1976) and China (Hou 1982).
Our objective in the comparative study was to
determine the respective petroleum potentials of
selected hoceue/Olpocene sequences from
Australia and India by using ostracode-based
parameters developed by Pokorny (1965) and
Oertili (1971) and tested recently by Giuhs (1983),
plus some other ostracode (Peypouquer 1979) and
sedimentary characteristics regarded by us as
pertinent.
it seemed to us that the most pragmatic test of
the relevanee of the study parameters would be to
determine them for wellsite samples. Nevertheless,
7 Riverina-Miunray Institiate of Higher Education, Wagna
Wagga, NSW 2650,
|) OW and Natural Gus Coomission, Bombay, 400078,
India,
outcrop samples,
including several collecred by us both on a brief
field excursion during October 1983 to Aldinga
Bay, South Australia, made il clear that outcrop
material would also yield consistent results, Of the
pluneering studies in this methodology that by
Pokorny (1965) was based mainly on outerap
samples but included specimens from two
boreholes, whereas Oertli (1971) worked exclusively
with wellsite samples. Im our study, Guha
determined the selected parameters for mne samples
from the Cambay Well, Cambay Basin, India,
while MeKengic deierniined them for the South
Australian Department of Mines and Knerpy
(SADMEB) Light | Well, in the Adelaide Plains Sub-
Basin, South Austrafias and also for the outcrop
samples.
‘Stratigraphic Summary and Material
South Australia
Ihe Adelaide Plains Sub-Basin is part of the St
Vincent Basin (Fig. 1, locality 1), MT hits been
extensively drilled to develop the groundwater
resources of the Adelaide region for which it
contains Iwo of the pritcipal wguilers.
Consequently, the subsurface stratigraphy is well
understood (Lindsay 1969, 1985). The Pulaeogene
sediments include both marine and continentil
sands, marine limestones and marls. They indicate
alternating shallow marine (inshore to outer shell),
transitional and fluvio-lacustrineg Palaeogene
palacoenvironments (Lindsay 1969, Cooper 1985:
Harris 1985}.
Wh kG. MeKENZID ADR. GUHA
Gul
St Vincent
Fig. 1. Locality map of South Australia indicating the
lucatbons of: 1, the SADME Light No. 1 Well (about
138" 26'F, Long, 34° 26'S. Lat.); and 2, the Maslin
Cay/Aldinga Ray coastal section of the Willunga
Embayment (hachured) of the St Vincent Gulf Basia
(dashed ourliney, Seale bar = 20 km. The general
Incation js arrowed on (he inset map of Australia,
The SADME. Light } Weil (reference bored, core
| of Lindsay 1969), is particularly noteworthy
heeause if contains, in a cored interval, the
Eocene/Oligocene boundary as estimated by
Lindsay (1969, 1985) near the base of dark grey,
cherty caleareous siltstones forming the "siliceous
unit” of Port Willunga Formation — the Ruwarune
Member of Cooper (1977, 1979), Below this He
equivalents of the latest Eocene, basal Aldinga
Member of Port Willunga Formation, comprising
ouleareous mudstone, greensand and siltstones grey-
brown speckled green, glauconilic, pyritic,
carbonaceuus, shelly; in part with very fine quartz
sand. Beneath a regressive pebbly sand correlated
with Chinaman Cully Formation (Lindsay 1985),
Fig, 2), an abbreviated interval equivalent to basal
Blatiche Point Formation (Lindsay 1968!, 1969)
resis.on, and fills. fractures jn probably Proterozoic
quartzite (Cornish 1964), Our material includes
seven samples supplied by J, M Lindsay (SADME)
' Lindsay, J. M. (1968) Palacontology and stratigraphy.
Appendix C, Vol. 1, /m Northern Adelaide Plains
groundwater study to May [968.2 vols, Depe of Mines
und Energy Repoet 67/123 (unpubl }
from the Eocene/Oligocene boundary zone i i his
well, i.e. From basal Ruwarung Member and
uppermost Aldinga Member of Port Willutiga
Formation.
Tetal depth of the SADME Tight | Well was
171.9 mi. The seven samples provided iit is froin
core 1 came from the following deprhs: 139.7 - 139.8
m; 139.8-140.) my; 14).1-141.2 ms 142.2-142,3 m;
142.3 -142.4 my 142.75-142.85 my and (43.2-143,3 m.
The Bocene/ Oligocene boundary as determined by
Lindsay. (1969) lies between I4].2 and [42,2 m in
this borehole,
Aspects of tte regional stratigraphy were
described in detail by Cooper (1979) in lis study
of the Willunga Eimbayment based en bores and
the classi coastal section [Fig. 1, locality 2),
Cooper (1979) concludes thar the Willunga
Embayment was a structurally contralled palneo-
bay for much of the Cainozoic. Sedimentation in
this embayment of the eastern St Vincent Basin
began in the Middle Eocene With {luvial sands and
intermittent carbonaceous swampy sediments
(North Maslin Sands), In the Adelaide Plains Sub-
Basin, where SADME Light | is located, lignitiv
Clinton Formation sediments were deposited next,
followed by the onset of marine transgression
(South Maslin Sands) involving reworking of rhe
earlier (inviatile sands as well as deposition af inter-
digitating marine and fluvio-lacustrine sandy
sediments. Further trimsgression was miarked by
high energy bioclastic limestones tich mm goethite
pellets (Tortachilla Limestane) overlain by
glauconitic, spicular and marly clays and silrs
(Blanche Point Formation) as the transgression
attained its maximum level during the Late Eovene,
Scasonal upwellings probably characterised this
interval. There followed a brief regression
(Chinaman Gully Formation) but then marine
conditions returned (Port Willunga Formations.
The sediments indicate inland to: coastal lateral
facies variations tram non-marine and marginal
sands ty richly fossiliferaus marine carbonates.
Such facies persisted from latest Eocene throughout
the Oligocene atid into the Miocene (Lindsay 1967,
1969, 1985; Cooper 1979), Qur material includes
four outcrop samples from the Eocene/Olivocene
boundary vone (Lindsay 1967; Lindsay &
McGowran 1986) in the coastal section at Aldings
Bay (Fig, |, locality 2).
India
The Cambay Basin (Fig. 2) has India’s largest
onshore oilfields and is ranked second alter the
? Goriish, B. E, (1964) Light No, | Well completion
report, Dept. uf Mines repare 39/007 (unpubl
OSTRACODA AS INBICATORS OF PETROLEUM POTENTIAL 7
CHING
~
Fined
PARISTAN 74
f
~ ~.
™ NEPAL ~~?
~_S
ambuy Well
Bomhay
Ny, 2. Locality map of India indicaring the Cambhay Basin
(hachured) and the lncution of the Cambay Well (72°
4o°b Long., 22° 22'N. Uat.), Seale bar - 400 ka,
Rombay offshore region in terms of reserves of
hydrocarbons (Guha 1983). Not surprisingly, tt has
been studied in considerable detail and lhe
Palaeoxene sequence is confidently correlated and
biozoned, based mainly on furaminifers.
Halaeoecological ssulyses indicale alternating
shallow marine (inner to outer shelf) and
transitional depositional environments for the
Basil during the Palaeogene (Guha & Singh 1980).
The Cambay Well is one of several thar were
drilled 16 about 2000-3000 m depth with the
primary objective of determining the basinal
lithostratigraphy.. lt bottomed in Late Cretaceous
basalts of the Deccan Trap. The Kocene/Oligocene
boundary in the Cambay area occurs in the larapur
Shale (Late Bovene-Oligocene) which in this well
overlies Cambay Shale (ltarly-Middle Bovene), The
intervening Vaso Formation (Middle-Late Eocene)
which is usually unfossiliferous was not identitied
in the Cambay Well (Guha 1 unpubl,),
The Tarapur Shale is variegated, grey-greenish
to fight brown, soft to fairly hard and poorly fissile.
I is characterised by thick intercalations of
quartzose,. fine-medium but occasionally coarse-
erained sandstone; and, in the Cambay area, by
thin intercafations of limestune near its base, The
underlying Cambay Shale is dark-colouree-
bituminous, moderately hard to fissile, with
occasional siltstone beds (Guha & Singh 1980),
Guha examined six samples from the Tarapur Shale
and three from the Cambay Shale,
Total depth of the Cambay Well was about 2500
m. The 9 samples available to us came from the
following deprhs: 1520 m; 1530 m; 1835 m; 1548: m;
$580 m; 1555 m (Tarapur Shale); and 1870 m;
L585 m; 1605 m (Cambay Shale). The
focene/Oligocene boundary ligs at about 1540 m
in this well,
Vhere is considerable support for an Oligocene
prospect (rom the offshore Cambay Basin, Guha
& Pandey (1980) in a study based mainly on the
Tarapur Ollshore Welland incorporating micro-
faunal, palynological, lithologic and electric log
analyses interpreted the wuerval wbuove the uis-
appearance of Hantkenina (Late Eocene foram-
iniferal datum) as beginning in deeper marine
basinal facies and proceeding upwards (above an
unconformity/disconformity) into two alternating
repressive — esiuarine and terrestnal — une
Transgressive — shallow marine — cycles befure the
uppearance of a characteristic Lower Miocene
assemblage. Except for the final (younger) shallow
Marine transgression there is good evidence for
abundant organic matter in euxinic deeper facies
and in shallow deltaic and paludal facies also
characterised by rapid sedimentation and burial
(Guha & Pandey 1980),
Methodology: Ostracode and Other Parameters
Carapaces/ Vuives ration
Use of this ratio to yield palacoccalogical
information was pioneered by Pokorny (1965) in
a wide-ranging paper which also dealt with the
implications of changing sex ratios and variations
in shell ornament. The changing sex ratio parameter
has not yet been turned to account for petroleum
exploration. On the other hand, many variations
in shell ornament are now interpreted in terms of
the rhopic factor (Peypouquet, ef.a/. 1982) and can
be used in suitable Facies Lo suggest presence or
ubsence of upwelling (McKenzie & Peypouquct
1984) which, in turn, is linked to the abundance
of organic matter.
Oertli (1971) reviewed Pokorny’s work and
telated the carapaces/valves ratio to potential lor
the formatien of hydrocarbons. Insummury, when
the ratio is high, rapid sedimentation — which
Minimises disarticulation of carapaces |Hto separate
valves — 1s indicated; and with sulficiently rapid
burial organi¢ matter is not absorbed by mincral
particles and so retains the potential for conversion
incu lydeovarbuns.
\8 Ki
Adults/ Juveniles ratio
Ii any inilerprelation of a palaeoenvironment
which is based on ovulispecies assemblaves it is
esserioal to separate the autochthonous fron. the
ullochthotious [anal elements. Ostracoda are one
of the groups in which Unis separation is achieved
rapidly because they molt trequently in progressing
lo adulthood and hecause juvenile shells, being alsa
calcareous, are usually preserved. Reyment {1971}
Uiscusses in derail ihe preparation of a life table
in order to understand the population dynamics of
species, We note that juvenile mortality is always
high in his examples based on gutochthonous.
Ostracoda, Therefare, the adults/juveniles ratio
should incicate a dominant percentage of juveniles
in autochthonous lossil populations, but be heavily
biassed towards either adults only ar juveniles only
for allochthonous taxa. Such biases are interpreted
mast reasonably as post moriem sorting efferts, OF
course, In some environments, <2. shorelines, post
mortem sorting Is charavteristic for all fossils,
autochthovons as well as allochthonous, Thus, to
be usefu) as a positive indicator of petroleum
potential, a relatively low adults/juveniles catio
feeds to be linked with a high carapaces/ valves
ratio, This is hecause the lovter indicates rapid
sedimentauon, which cends 10 minimise sorting,
Percentage of fragcmens
This useful parameter 8 usually ignored by
workers making up assemblages slides who tend to
pick whole spevimiens. The percentage of fragments
is simple ta obtain: two or three Counts of 106
specimens, inclusive of all fragmentary ones, bein
sullicient to estimate it reasonably for any washed
sample, Obviously, care must be exercised to avoid
damaging specimens during preparation al’
washings; and the parameter cannot be used
conveniently, with indurated sediments,
When there t} a Significant percentage of
fragments. it implies a high energy enyireninenr.
Low pervenriges inclicate law energy environments
or else rapid sedimenation,
Crushed or warn specimens
Commonsense sligrests that, where
autochthonous tava are concerned, worn or
abraded specimens aré indicatlve of hieh energy
environments and slow sedimenation, Far
allochihorns, abrasion is another paramerer by
which their allochthonous pravenance can be
interpreted — in the case where autochthonous
species are well preserved (nol abraded). On the
other Ward, crushed specimens are assoeiated
typically with line grained muddy and marty
offshore sediments. When abundant they ate
interpreted as indicating considerable compaction
MeoRENZTE & 1. KR. OUTTA
pressure, as might be caused, for example, by rapid
olfshore sedimentation,
Krithe and Purukrithe
The podocopid ostracode genera Arie and
Parakrithe are used as palaeoecologie Indives by
Peypouquer (1977, 1979) and others to interpret (he
palacohydrology of marine sediments, in particular
palaeodepths but alsa dissolved oxygen (O.}
content, lool supply tnutrient) and upwellings.
Peypouget’s hypothesis is that the size of the
vestibule in the non-calcilied inner laniclla in Arise
and Parekrisie is more or less inversely
proportional to the dissolved 5 content of the
ambient seawater.
Peypouquet (1977) proposed a physiological
explanation based on the known inverse
relationship in crustaceans between external
dissolved O and organism haemozlobin (HB}, In
uw eurrent review, McKetiele (1986) considered the
effects upon MB of several ather enviranmental
actors and found that those which were significant
— PH. food supply and subscrate lerrows iron —
would all affect HB synthesis in the same direction
as dissolved O., This result reinforces the
hypothesis, Reeenly Aladin (1983, 1984) used
microcryoscopical techniques ita show dio-
chemically that HE regulation in puducopid
Ostracoda oceurs probubly via salt sequestration
in the noncalcified membraneous part of rhe inner
lamella. This is precisely the site which Peypouguel
(1977, 1979) hypothesised would reflect HB
respunse by Avithe and Parakrithe to dissolved Os
variations, The hypothesis has been upplicd
frutt Cully in interpreting several palacoenvironments
ranging in age from Manstrichtian to Miocene
(Donze et al. 1982; Peypouquet er af 1982;
Mckenzie & Peypouquet 19844,
Pyrite wind Gypsum
In Oceatic sediments, sulphur is about cqually
divided between salphare and sulphide species. The
lractionation from sulphate cto sulphide
(pyrilisation) is due ro desulfobacterial sulphate
reduvtion, The oecurrence of pyritisation in a
palacocnyironment i§ readily interpreted by
reference to its microfauna. Vor Ostravoda.
pyritisalion is indivated by a signifient percentage
of brownish to blackish valves and carapaces
Oceasional spevimens alisten with pyrite whieh ts
the diagenetic oxidised sity of the pnmary reduced
ferrous sulpfide that stains/permeates shells to
produce the chagnostic coloration
Cypsurn is the predominant form of sulphate in
marine sediments where it is a component in a
ulohal carhon/sulphur redox system that
incorporutes carbonates, sulphates, sulphides amd
OSTRACODA AS INDICATORS OF PEI ROLEUM POTENTIAL, i9
organ carbon (Carrels & Lerman 1984). In
weathered Gutcrops, its presence is signalled by
chimps of relatively large and often Fragile evpsum
erystals; fut in unweathered sediments it can occur
4S (presumably diagenetic) microspherulites. ‘The
laller were recorded, for cxample, in the Mioecne
Fyansford Formation, collected ar Balcombe Bay,
near Mornington, Wietovia ([McKenaie &
Peypouquet 1984); pypsuin crystals are common
ia Weathered oulcrops at the same localiry.
When mirine sedimentary sequences carry
abundant sulphate/sulphide. their pdlaco-
citvironment 4s imterpreted as having been
Ucoxygenated and strongly reducing in the presence
of ubundant decaying organic matter and
imlerstitial plus seawater sulphate. This
interpretation derives from their sulphide content,
Even when diagenesis and weathering have led
tainly to Yormation by oxidative processes af the
sulphate pypauni, the slammed shells of the
microfauna are palimpsests of a depasicional
reducing palaeoenvironment. A reducing
cuvitonment, churdacrerised also by rapid
sedimentiuton/ burial, inhibits oxidation ol urganic
matter hydrocarbons and, under appropriate depth
ef Pdnal and temperature conditions, yields
petroleum,
Cilaucontte
Glauconite is an easily pecounisable green iron-
bearing Silicate which 1 associared wilh stable ourer
shel! environments, slow sedimentation,
moderately anaerobic conditians on the bottom,
and 4 large amount of deeaying organic matter.
Ollen, it decurs together with pyrite. Its significance
in source rock ierpretanan siems from the
association with slow, even negative, secimentation
because under such conditions organic matter
hecomes oxidised and is no longer available to
generate hydrocaibons. Glauconite praing are
readily transported and sorted commonly forming
greensands, Such deposits, with the inpligation of
bollom current action jn addition to slow
sédimenation, alo are cuunlerandicators in
petroleum source Tork interpretation,
Results
ables ) und 2 provide results of determinations
of several of the parameters discussed ubove for
samples from the SADME Light | Well, Adelaide
Plains Sub-basin. Sourh Australia, and the Cambay
Well, Cambay Basin, India, respectively. "There
were no significant occurrences of erushed or
abrailed Specimens in the counts we made, and
eypseous tnicrospherulites were only searched for
in the South Australian samples, in which they were
uniformly rare.
The results indicate: that similar numbers of
specimens were counted for both sets of samples:
that the carapace/Valves ratin is constantly higher
in the Cambay Well as is pyritisation; that che
percentave of fragments’ is much grearer and that
elaucanite is only abundant in SADME Light |;
that the percentage of adalts is similar in both
sections — varying trom 14.5-45% (mean 265%)
in SADME Light 1, and from 15-44% (mean
33.4%) in the Cambay Well.
The Jast statistics suggest that we are dealing with
assemblages which have approximately similar
population dynamics. and are predominantly
autochthonous (both average about 70% juveniles).
Thus, allhough the two wells are widely separated
geovraphically, {hey may fairly be compared using
ostracodes as indices for source rock characteristics.
The combination of high percentages of
curapages and pyritisation in the Cambay Well
section especially in the lower Tarapur Shale and
in the Cambay Shule (Table 2) suggests rapid
sedimentation offshore combined with a reducing
environment in the presence of abundant decaying
Orgame mutter. As organic matter decays it uses
up available oxygen. This is confirmed by the
resulis [ram {he occasional valves of Krithe and
Perakrithe in the Cambay Well assemblages. In ihe
upper Tarapur Shale samples, whove the
Eucene/Oligecene boundary, the Kritke have small
TAM 1. Some parameters of Ostracoda in SADME Light 1 Well, Adelyide Plains Sub-basin, South Australia;
recoded in percent excel for tand & a — abundanize = common, | = carapaces; 2 =
4 whauconite; 5 = pyritisarian, 6
adults; 3 -— fragments;
turnbers of specimens ((he figures i parentheses are the numbers af specimens
excluding fragments}, he Fovene/Olizacene boundary is indicated ar the base of the Ruwaruny Member, Port Willunga
Formation (PW),
t 2
Kuwarung Member, PWF 4.6 M1
Rirwarung Member, PWE 5.4 14,35
busal Ruwaruny U6 22.4
yop Aldina Member, PWEF 3.6 26.2
lop Aldinga Member, PWF 21 241
lop Aldinga Member. PWE 7 m4
tup Aldinga Member, PWR 250 45)
.
4 4 5 fi
45.1 a 12.0 217: (118)
30.0 a a5 410217)
WI a 15 271 (169)
55.7 4 4.0 305 (135)
53.4 a 15 285 (133)
19.8 a 10 244 (147)
20.0 © nA 40 (42)
—_—_—— Oe OO ees
mt) K.G. McKENZIE & D. kK. GUHA
TABLE 2. Some parantvters af Ostravoda th the Cambay
Well, Cumbay Basin, [ndia, recorded im percent except for
4und 6» — rare, SLR. — noi recorded. | = carapaces;
2 = adulis, 3 = fragmeénisy 4 = glauconites 3 -
pyritisarion; 6 nombers of specimens (inetoding
Irizments). The Bocene/Oligocene boundary 14 ingticuted
in the Turupur Shale.
| 2 3 4 5 h
Tarapur Sluale 52. 30 7 v 32. (156
Tavapur Shale 49° 38 13 t 37 s«78
Tarapur Shale Ss} 43 #11 c 230 ON
Tarapur Shale 72 37 #612 f 6& 24
farapur Shale 9 26 1 NLR 5 196
Tarapur Shale = $2, 1S SCNT 276
Cambay Shale 100 4t 42 NLR. O82 115
Cambay Shale 100 44 28 WLR 87 132
Cambay Shale 100 29 7 NLR 9S RS
vestibules indicative of a well oxygenated milieu:
in the Jower Tarapur Shale sampies the Arirfe have
very large vestibules which indigales an oxygen poor
tur reducing) environment. Untortunalely, no Krithe
or Parakrithe were identified in the three Carbay
Shale samples,
On the other hand, inihe SADME Light 1 Well
avction (Table 1) rhe percentage of carapaces is low
in all samples (2.1-9.6%)) except the oldese (254%)
ald the percentage of fragments is seniticuntly high
(30,0-55.7%) except in the oldest sample (20%),
Further, glauconite is abundant in all samples {it
is tare OF not. recorded in the Camhay Well} except
the oldest sample, where it is common; and
pyritisation is abyays rare except in the uppermost
sample. These dala suggest thal marine
sedimentation in the SADME Light | Well area
during Egeene/Oligocene boundary Lime twuk place
ora well oxygenated outer shell characterised by
slow sedimentation; and relatively constant bottom
traction — Which produced the large numbers of
shell fragments. The occasional Arithe which occur
in these assemblages confirm the well oxygenated
orlicu since they fave small yestibules.
We conclude that, in lerms of the ustracude and
other parameters we have siudied, the
Loone/(Nigocene boundary vone sedinjents lave
good petroleum potential in India but lil!le potential
in South Australia. This conclusion js consistent
with che drilling resiulis. The Adelaide Plains Sub-
basin has been extensively drilled as part of a
thorough aquifer explorauon programme without
Providing any satistactary indications of petroleum
hydrocarbons Jo [ndia, the Cambay Basin is second
ody to the Bombay Of shore Basin in hydrocarbon
reserves itl hits Enclia’s largest onshore oilfields.
The source racks are located in offshore facies
immediately below the bocene/Oligocune boundary
and extending downwards into the Middle-Rarly
Facene sediments,
Detailed Analysis — South Ausrralia
Table 3 provides. the results ol a nare detailed
analysis of the SADME Light 1 Well samples, This
analysis intioduces another paranicter, Ihe number
of ostracodes per gram of washings pickeal, as well
as breaking down the assemblages imto lamilies and
analysing these both compositionally and in terms
of the patametets alreudy studied lor the
assemblages as a Whole. The prime objective is to
enable a more precise palaedecological
interpretation,
All the washings were weighed and then picked
for their ostracodes. In some cases, the entire
fraction had to be picked to yield a satisfactory
count fabeut 100-200 specimens) but in ome
instance (Ihe stratizraphically lowest sample) even
this method yielded only 40 specimens, including
fragments. In most samples, however, Ostracoda
were sO abundant that only a fraction of the
washings teeded to be picked, This was (he case
especially with the two lowest samples from the
Ruwarung Member of the Porte Willunga
Formation. ‘The richest sample was the middle
sample of this Ruwarung Member series and the
poores! was the lowest Aldinga Member sample,
Ti terms oF ostracude diversity, most of ihe
samples seem very similar, having 19-20 species; but
the two lowest {Aldinga Member) samples are Less
diverse carrying 15 and 1 species respectively, Since
diversity can inctease significantly with higher
counts (Cronin 1984; Whatley & Dowoing 1483)
these diversity results do tot warrant more derailed
comment.
Tatlt 3. Anelysiy of oxtrocode relative abundances in
selected faninles for the SA DME Light | Well frenresentiag
93, 7-99.2% wf Jotal ustracodes in the samples examined)
Duta reverted in percent. Also recorded ure the
astracodes‘em data. AG samples examined (a5 lar
‘lable 1). with A | uppermost Rowarung Member saraple
ta G = lowest Aldings Member sample, respectively.
Ortracode
Family A BR «€. TR Ber F
Bythocytheridae 1.8 OO 4,1 9S 10.2 OF 5.0
Cytheruridae 14.8 97 TO UR Th 9.0 12S
Xestoleberididae = =4.2 12.5 100 3,3 3.2 45 10.0
Travhyleberididae 46.1 3L.9 $5 4 43.9 $1.9 58.6 15.0
Krithidae On 2.3 3.0 0.3 0.0 O4 CO
Pontocyprididae 83 23.9 92 11,5 105 6.2 %hO
Paracypodidae 00 $5 85 00 00 OO Oa
Macrocyprididae 0.0 0.0 1.7 Of) WO O4F oo
Bythoxyididae 0.5 2.6 0.0 0.0 0.0 0.0 0.40
Cytherellidae Ib 9.7 LLL 17-4 10.9 19.7 S.0
% of total
ostracndes YT YT YU) 94 84 992 47S
Qther Parveietery
Osiracsodes/om (less fragments)
14.9 15 52.6 15 169 186 20
Ostragndes“am finvhidiny Cravens)
25.6 LSS) $4.7 43.6 36.5 298 15
OSTRACODA AS INDICATORS OF PETROLEUM POTENTIAL, 2l
Wherr lhe assemblages are broke: down into their
component families it seems clear that the two
Vichese saniples (Ruwarung Member) represent an
offshore, even Outer shelf environment. Both carry
several specimens (7-8) of the genus Arie which
i chagnaste lor uuler shelf and deeper waters.
Additionally, the middle Ruwarung Member saniple
(8B, in Table 3) has the family Bythocyprididae
(genus Bythocypris) which also is typical of deep
water facies. This fauna occurs a little higher than
the Eovene/Oligocenc boundary as determined on
foraminifera (Lindsay 1969) thus its
palagoecological interpretation is consistent
stratignaphically with what we know of ostravode
trends worldwide (Benson 1975) and with the data
from Ostracoda of the Willunga Embaymenr
(McKenzie, in Cooper [979t.
The dominant familes are the Trachyleberididae.
Cythecuridae Xestuleberididwe, Pontucyprididae
and Cyiherellidae. Several other familiees are
represented by so few specimens that they have not
been included tn Table 3, which lists only the Tu
inust abundant families — representing 93,70 to
949.2%) of total ostracode assemblages. These poorly
represented taniilies include J oxevenchidlae,
Cytheridae (genus Loxocythere), Eucytheridae
(yenus Rotundracythere), Leplocytheridac,
Schizovytheridae (genus Patjenborchella).
Surprisingly, no specimens of Bairdiidae were
identilied although this family as virtually
ubiquitous in marine facies. The absence of
Kairdlidac seems ta be a local variation because the
family cerlainiy occurs in coeval sediments [mom the
Willunga Embayment (McKenzie 1979).
Clearly, the mov abundant family is the
Traclvleberididae (Table 3); although 1 is Jess
abundant (31.9%) in the richest sample (B) than in
the arher sainples (45-58.6%). On thiy ground, we
decided i study trachyleheridid data for the
Tsniv 4, Sune perameters ef Trachyleberididae in [he
S4ADME Light 1 Well, Deta recorded in perveni except
column 5S which gives actuel numbers of specimens
analysed, \ = carapaces; 2 = adults; 3 = tragments;
4 - pyrilisation. Thus, for the Ruwarung Member. PWEF
there are: 2% carapaces, (98% valves}; 30% adults (70%
juveniles); 57%) fragments: and a quarter of the specimens
are pyriused.
L 2 3 4 4
Ruwarung Mewber, PWE 27 W i7 25 100
Ruwarung Member, PWF 5 68 40 10 99
basal Ruwarung Member 10 30 40 5 150
top Aldinga Member, PWF 3 30 63 7 134
Aldinga Member, PWF a 22 63,5 6 148
Aldinga Member [WEF 7 25 40 25 \43
Aldinga Member PWT 2 33 O33 0 18
parameters which we used in the preceding more
penenal enalveis (‘fable 4).
We infer, plausibly, that ia the uppernsosi
Ruwarung Member sample bottom conditions were
reducing for at least part of the time. Vhis suggests
that elsewhere in the Becene/Oligacene of Suuth
Australia the Early Oligocene might be a
prospecove petrolilerous zone if the sediments are
thicker and also contain high numbers of carapaces,
fewer Traginents and much less glauconite,
‘The Encene/Oligocene boundary zonets already
a drilling target off Gippsland, Victoria (Douglas
& Ferguson 1976) but with respeet to much thicker
sections than occur in the St Vincent Basin.
Unfortunately, our results from SADMF. Light 1
offer only slight encouragement for a more intensive
exploration of this interval in South Australia
Compurisor with Oulerops
During October 1983, we sampled the classic
coastal Eocene/Oligocene sections at Maslin Bay
and Aldinga Bay, South Australia. Table 5 provides
a resume of the data on ostracode parameters and
glauconite for samples from this collection.
Although the SADME Light | Well (Table 1) was
much more closcly sampled (over only 1.4 m in the
basal Ruwarung Member and only 22 m in the
upper Aldingu Member), the ostracode data [rom
outcrops correspond rather well. Ln particular, the
Percentages of carapaces for the two lawest Aldinga
Member samples of SADME Ligh | {located abuul
1.6-2.0 m below the basal Ruwarung Member
sample) are sinwilar in tle two outcrop samples trom
the Aldinga Member (Table §), Generally, the
outcrop samples are less pyeitised.
Conclusians
Ostracode parameters, reinforced with evidence
from pyrite, gypsum and glauconite can be used tu
indicate petroleum source rock potential in the
enclosing sediments,
lL. A high percentage of carapaces indicates rapid
burial,
2. Large percentages of adults and juveniles (with
juveniles. dominant) indicates a mostly
aittachthorimus community,
_ A low percentage of fragments indicates low
cnerey and minimal bottom cucrents/traction,
+. A high percentage of pyrilisation of ustracode
earapaces and valves indicates a reducing
environment; as do diageneti¢ pyrite and gypsum
in the enclosing sediments,
These several characteristics all inlieate oud
petroleum source rock potential as our extended
discussion has made clears.
'
Le
22 &G. McKENZIE & DK, GUILA
LAME 5, Some purunielers of Ostracoda, plus alaucanife in ogtcreps uf Aldinga Bay, Sauth Australla, recorded
jn percent except far 5. t — trace; | = carapaces; 2 =
adults; 3
fragmenta; + ghuuyanites 3 = pyritisations
6 — number of specimens ((he figures in parentheses are the mumbers of specimens excluding fragments). The
Eovene Oligocene boundary is indicated.
1
3m above hase of Ruwatung Member 1.4
MY om below top of Aldinga Member a9
4 nt above base of Aldingau Member 25.4)
2m below top of Gull Rack Member 124
2 3 4 5 6
AW.2 52.8 1 nil 237 (112)
25.8 50,9 12.5 l 236 (1161
25.0 Us t mil 164 (96)
is. 24.6 MW) fit 225 (168)
Inthe exemplar series, astracode parameters and
high pyritisation all confirm the high petroleum
source rock potential of the Eucene/Oligocene
houndary zone in the Cambay Well, India. On the
other hand, in the SADME Ligtt ¢ Well, South
Australia, and. in outcrop samples from South
Australis the only positive correlations are with the
dominantly autochthonous community
characteristic (2, above), Im other respeets, the
Australian samples of the Eucene/Olizecene
boundary zone correlate negatively with indications
of good petraleum source rock potential, Likewise,
elyuconite, which fs counter-indicative of
petroleum source ruck potential, i5 generally
common to abundant in the Austrahban exemplar
samples but is not recorded or rare i the Cambay
Well.
IL is not Surprising, therefore, that the Catmbay
Basin is India’s major onshore oilfield in
bocene/Olipoceneé strata whereas South Australia
is non-praductive for this interval in the section
concerned. Gur present methodology appears to be
ay CHective for the Eocenc/Oligocene of Iidia and
Australia in the areas tested as similur bul less
detailed methods have proved previously in the
Turonian-Coniacian of Bohemia (Pokorny 1965),
the Neucomian-Aptian of southwestern Fraice
(Oerth 1971), the Maastrichtian-Palaeovene oF
Mozambique (Qertl 1971) aud Bathonian-Tertiary
uf India (Guha 1983),
Finally, when these parameters ate reassessed Por
samples iu which the Ostracoda huve been divided
imo their component families as done (K.G.M.) for
she SADME Light | Well, the results alluw an
opportunity to achieve a consisient but more
detailed palaeoecological interpretation than could
result from the gross data alone.
Acknowledgments
D. K. Guha acknowledges with gratitude support
Hon the Australia/ India Scienve and Technology
Agreement enabling his visit to Riverina-Murray
Institute of Higher Kducationd{R.M.H.B.) where
the joint project was carried out, Both authors are
grateful to Mr J. M. Linday and Dy B. I. Cooper,
South Australian Department ef Mines and Energy
(SADME), and to Dr B. MeGowran, Depactrrent
of Geology and Geophysics, the University of
Adelaide for guiding them throurh the classic
Tertiary sectians at Maslin Bay and Aldinga Bay,
neat Port Wilhinga, South Ausiralia, Mr J, M.
Lindsay is also thanked for loaning the samples
from SADME Light 1 Well, and lor his con-
structive comments on the paper. Professor 1, P
Peypouquet, Université de Bordeaux, ig thanked
for a pertinent review. K, G. McKenzie
acknowledges ARGS Grant No, E80 15387, DR.
Guha publishes with permisston from the General
Manager, Oil and Natural Gas Commission, India.
Some Tables and the tWo Figures for this paper
were ¢xhibited at the 8th Australian Geological
Congress, held at Flinders University, Bedford
Park, South Australia, during February 1986.
Kk. G. Mckenzie acknowledges support from the
R.M.NHLE. Staff Develapment Prograuuie
enabling his attendanee at the Congress. Mrs Jan
Scaman um! Ms Colleen Seberry typed the
Manuscript,
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THE CLASSIFICATION OF THE MITE FAMILIES TROMBELLIDAE AND
JOHNSTONIANIDAE AND RELATED GROUPS, WITH THE
DESCRIPTION OF A NEW LARVA (ACARINA: TROMBELLIDAE:
NOTHROTROMBIDIUM) FROM NORTH AMERICA
BY R. V. SOUTHCOTT*
Summary
Amongst the Trombidioidea an unnamed family group containing Trombellidae, Chyzeriidae and
Audyanidae Fam. nov. is recognized; these families are defined and keys provided for the larvae of
the families, subfamilies and genera. The Johnstonianidae is examined, and three new subfamilies,
Tetrathrombiinae, Pteridopodinae and Ralphaudyninae are established, with Ralphaudyna
Vercammen-Grandjean ef al., 1974 being transferred to the Johnstonianidae.
Ralphaudyna amamiensis Vercammen-Grandjean, Kumada, Newell, Robaux & Suzuki is recorded
from a second Japanese location, as an ectoparasite on the gryllacridoid Tachycines robustus Ander
(Orthoptera, Rhaphidophoridae). Further metric and descriptive data are given for this larval mite.
Nothrotrombidium treati sp. nov., larva (Acarina: Trombellidae) is described from a single
specimen found dead on a noctuid moth Spaelotis clandestina (Harris) (Lepidoptera: Noctuidae) at
Tyringham, Mass., U.S.A.
This is the first record of this genus in North America, previously recorded from Europe and South
America, as well as Madeira, in the Atlantic Ocean, and Asia.
KEY WORDS: Taxonomy, Nothrotrombidium, Ralphaudyna, larva, North America, Japan,
Acarina, Trombidioidea.
TITE CLASSIFICATION OF THE MITE FAMILIES TROMBELLIDAE AND
JOHNSTONIANIDAE AND RELATED GROUPS, WITH THE DESCRIPTION OF A NEW
LARVA (ACARINA: TROMBELLIDAE: NOTHROTROMBIDIUM)
FROM NORTH AMERICA
by R, V. SOUTHCOTT*
Summary
SouTHeart, R. V, (1987) The classification of the mite families Trombellidae and Johnstonianidae and
related groups, with (he description of 4 tlew farva (Avarima: Trombellidaes Vothratrambidiin) trom
Norrh America. frams. A. Soe. 8. Aust. WAL), 25-42, 29 May. 1987.
Amongst the Thombidioidea an unnamed family group containing Trombellidae, Chyveriidae and
Andyanidae fam, mov. 1§ recognized: these families are defined and keys provided for the larvae of the lamiilies,
sublanntics and genera, The Johnsionianidae is-examined, and three new sublamilies, Terrarhrombiinie,
Pleridopodinae and Ralphaudyninae are established, with Ra/phaudyad Vercummen-Grandjean eral, 1974
being transferred (o the Johnstonianidae.
Ralpheudyne amamiensis Vercammen-Grandjean, Kumada, Newell, Rabaux & Suzuki iw recorded from
asecond Japanese location, as an ectoparasite on the gryNacridoid Jachyeines robustas Ander (Orthoptera,
Khaphidophoridae), Vurther metric and descriptive data are given for this larval mute,
Nathrarrombidtiar treatt sp. nov,, larva (Acarina: Trombellidae) ig deseribed from a single specimen
found dead on a Noctuid moth Spaélotisclandestina (Harris) (Lepidoptera: Noctuidae) at Tyringham, Mass,
USA,
This is the first record of this genus in North America, previously recorded from Furope and South
America, as well.as Madeira, in the Atlantic Ocean, and Asia.
Kry Worns: Taxonomy, Nethrotrombidium, Relphandyna, larvae, North America, dapam Acatrina,
Trombidiaidtea,
Introduction
The Johnstonianidae and the Trombellidae are
acvepted by some workers a5 the most primitive
families of the Trombidioidew. In the case of the
Johnstonianidac this opinion is based on che
presence of a number of supposedly primitive
morphological, hehaviqural and ecological
characters (Newell L957) Thus there are two pairs
of sensilla on the seutum, crista or equivalent areas
in most species, resembling the situation im the
ierythracvidea, but with loss or modification of the
anterior pair in some genera. Other supposedly
primitive characters are the simplicity of body setae,
the presence of larger than usual numbers of
sensory setae on various leg segments, and the
eeneral lack of reduction of leg segmentation, as
well as the presence of vaTious specialized setae on
ute gnathosoma, Some larvae are apparently
predatory, while others are, as in the majority of
the trombidioids, ectopurasitic upon arthropods,
Newell (1957) has also stressed (he “self{-detaching"
character of the larvae in response to stimuli, for
example when the host with its larval ccroparasites
is immersed in preserving fluids. There is also,
among the adult as wel as the larval jobnstonianids,
a sensitivity to heat and desiccation (Newell 1957,
196()).
© 2 Taylors Road, Miletwm, 5, Aust, 5062.
The Trombellidae (which have only one pair of
prodorsal sensilla) possess a number of simula
characters. The adult leg segmentation is not
reduced, and in the Jarvae the legs usually carry
more sensory setac than other trombidioids. The
larval coxue (end to he separated, and the urstigma
may not be strangly associated with coxa 1. The
concept of the Trombellidac as primitive may he
based more un a general resemblance to the
Johnstonianidae than on more specific indieatinns
Variant views are expressed by other workers
Thus Feider (1959b) eave an illustration
hypothesizing that the “Tanaupuslidae” and
“Calothrombiidae” ure che two most primitive
families of the Trombidioidea, originating [ron a
commen branch, while the next branch, ata slightly
higher level, gave rise to. the Johnstonisnidae and
“Notothrombiidae”™, On che other hand,
Vercamren-Grandjean ef a/. (1973) considered thie
the families Trombiculidse and Leenwentioekiidae
are more primitive than other “close families, such
us (he Johostonianidae ._ "4,
Thus the phylogeny of these mites remains a
matter of contention (an equivalent situation occurs
in the Drythraeoidea — see Southcott 196]a).
Thor (1935) divided the family Trombidiidae
Leach, 1815 (Trombidioidea of this paper) into 10
subfamilies, many of which have later heen
accorded family status Among such were [he
Johustomianinac and the Trambellinac, the latter
26 R. 8 SOUTHCOTT
delined a5 follows: Body clongale. Abdomen
rectangulat, Skin hardstitm ("hart"), rough-‘uneven
(“hockengz"), hairs short, pointed, Crista lacking:
the two sensory hairs sit close together in the mid-
dle of the horas in two thick tubercles between the
uwo scsuile eye pairs. The fourth palpal segment with
dilfering spines or hairs; fifth seament long,
Only the type genus Prompbella Betlese, 187, is
included {Iransfation RNS), There was no referenve
to the Jarvae. as then unknown. The adulr cenus
CéApzeru Canestnni, 1897 was omitted
Feider (1958b) deseribed the larva of the
trombythu mite Netkrotrombidium otiorvue
(Berlese, 1902), stating that this was the firsi genius
in the tamily for which there was correlation
between larva and adult. However, two genera,
Chyzeria sce Womersley. 934) and Avdvane (see
Wamenley [9$da,b), had previously been correlated
between the larva and the adult or deuronyinph.
These rearings allowed sunie attempts to define the
characters of the larval Trombellida¢ (Vercamimen-
Cirandjean ey af. 1974; Southcoart 1982). Following
avorrelation of an Apstalian Tanbelle larva with
its deulonvuiph, the classification of the
Trombellidae has been developed further (Southcott
[9B6u).
Nathrotrombidiiumn was fourded by Womersley.
(L9S4b), with type species Tramthella atorven
Rerlese, 1902 from Europe. In the genus he placed
alo 7) nothroides Berlese, 1888 from Sauth
America, and 7 lundhladi Willmann, £939, from
Madeira. A further species, No irevirersien Andre,
1960, hus been described [rom Nha Trang,
Indochina. All of these were adults,
In this pauper a secand larval species of
Nothrotrambidium, N. treari sp. noy., is described
from a nectuid moth in North America. This
disovery promprs a further examination of ihe
characters of the Trombellidac and related
trombidioid mites.
TASOQNOMIC DECISIONS AND ACCOUNTS
In the most recent reviews of the Trantbellidie.
Suurheott (1982, 19864) included the follawing
genera: Trombella, Chyzeria Canestrini, 897.
Bonierslevia Radlord, 1946, Nothomombicula
Dumbleton, 1947, Anvdvana Womerslev, 1954,
Nathrotrombidiunt Womersley, 1954, Durenia
Vercammen-CGirandjean, 1955, Parathrampbello
Andrée, 1958, Neanofhrothrombidium Robaux.
L968, Kalphaudyna Vercammen-Grandjean et a/,,
1974 aml Maiputrombella Southcott, LRG. OF these
Trembella, Chvzeria, Durenia, Axdvana and
Nathrothrembidiam ate knowir both as larvae and
adults or deutonymphs; MWorrersferia,
Nalhotronbicula and Ralnhaudvny are Kiown only
as larvae. and Puruthrombeltta,
Nearwhruthromivaiun and Malpotrombela ave
known only as adults,
Southcot (96a) excluded Paruchvseria Hitst,
1926, from the Tromibellidae, and plaved ti in the
Johnstoqiinicdac,
Vervaminen-Grandjean (1973) placed six
subfamilies to the Trombellidae: Trombellinac,
Tanaupodinac, Caluihrombiinae, Spelaeothrom-
bilnae, Noto! hrombiinae and Moyanellinac, He gave
no reasons for these decisions, which may have
stemmed largely fram the difficulties of placing
groups with which he was relatively unfamiliar. A
well-marked crista is present in the adults of the
Tanaupodinae, Calothrombiinae and Spelaeo-
thrombiinac, so there appears no reason to associate
them with the ‘Trombetlidac, Chyceriidae and
Audyanidae. In Nerothrombiunt Storkan, 934
(ihe sole genus of the Notothrambiinae) and In
Movanella Boshell & Kerr, 1942 (the sole genus of
the Moyanellinae} the crisia of the adults is poorly
defined or absenl; both have two pairs of prodorsal
sensilla (see Thor & Willman 1947; Rabais 1967);
thus it appears that their affinities le more with
the Johnstonianidae rather than the Trombellidae,
and they are here considered as being part al the
johnstonianld family group, possibly deserving
Tamily status. As the larva of neither of these two
families Js known, however, they will not be
comsidered further here
A difficulty in the classification af the
Trombidioidea (and other Parasilengona) is caused
by (he extreme heteromoarphy which exiscs beoween
the larvac and the adults (or deutauvinphsy), causing
dual generic und specific names for the hexapod and
octoped instars, as Well.as difficulties [a laaonomic
placements, failing accurate correlations, One such
instance is as follows,
Ralphaudyne Vereammen-Grandjean ef al, 174
was placed by its authors in the tribe Chyzeriini of
the Trambellinae |=lrombellidae of this paper). The
genus Was ereyied lor a single specimen of &,
ananienss Vercommen-Grandican ef al, 1974,
obtained in “soil under an-olden tree eave on the
middle slope of Mt Yawarn-dake Amant).
oshimea istind", Japan, and henee fren an
unknown host, or pocential host, arthropod. They
laid stress upon the “presensillac” Of the dorsul
idiasomal scutum, and therefore considered it a link
herween “the Iwo groups Chyzeria and
Parachyzeria”. However. the larva of Purdehyerta
has not been described. The placing of
Ralphaudyna in the Trombellinae was accepted by
Southicon (1982, [aséa).
Further specimens of Ratyhavedvna annuinien sis
have been Joutid al a sceond locality in Japan, taken
parusile ona eryllacrdoid (Opthoplera} see pr. 38),
MUTE CLASSIFICATION WW
Examination of these specimens stows that the
“nresensillae” or “presensilla” are close to typical
seutal (richobothria, placed more posteriorly an the
seituin, although smaller, The only passibly
Wuportant dilference between the anterior and
posterior trichobothria lies in the antetor ones
having & semewhul thickened shalt for the whole
of the sensillury seta, Ralphaudyna was placed in
the Chyzenini of the Trombellinae by tr authors
father than an the Johnstonianidae principally on
this character, and because it is “provided with w
very wide nasus. Several nude genualac on euch
low."
As fir as the preschee of “prosensilla” is
eoucerned, Newell (1958) used rhis term as
“prosensillar setav” but withour definitions in 1960
he defined aienn “prosensilliim” to apply to paired
ameromedian scutal setae of Trombiculidac,
Johnstonianidac and Trombidiidae, The presence
of such Setue, if ane accepts Newell's concept, is not
a detinilive character for any particular member of
the three tamilics nominated. Newell in fact stated
(YAO) (thar the term “prosensillum™” could be applied
in many Trombiculidae to the wapaired
allrerornedian scural set,
A prominent, even wide, nasus had been
described in various larvae of the Johnstonianidae,
eg. in Diplothrombiurn Berlese, 10 by Newell
(1957), Veider (19894), in Cearrorrombidium
Krumer, 1896 by Newell (1957), and in other genera,
Accepting the term “genuala” as applying to any
nude sensory sete on the leg genu (and excluding
the vestigiogenualie) such setae are absent in
Chyzerig and Nothotrombicula, but are present in
Trombella and Nothrotrombidium (Vercammen-
Cirandjean 1972; Southcort 1982, |9R6u) as well as
in dhe johnstonianid genera Lassenia Newell, 1957.
Oiplothrambium and others (see Newell 1957).
None of the criteria advanved by Vercammen-
Grandjean ef af (1974) for the exclusion of
Ralphaudyna trom the Johnstonianidae and. its
placement in the Trombellinae (Trambellidae) is
sustained,
Thus, Relghuudyne is temoved from the
Trombellidae, and placed in the lohristoniunida,
in Ralphiudyninag, subfam, mov,
Kranz (1978), following advice [i Wh.) from
VYercummen-Grandjvan, Has used the family term
Chyzeriidue (as Chyzeridac), separating it by key
characters’ from Trombellidae. Here | define the
Chyzeridac and a restricied family Trombellidue
Seta and seutal terminology follows Southoot|
(196)a,b, 1963, 1986a,h),
Superfamily Trombidioides tench
Monn! sereanwiny
lrombidides Leach J61S, pp. 387, 395.
Trombidiidae Michael 1884, pp. 4, 38 tad p.)s Thor
& Wilimann 1947, pr 187,
Trombidiinae Michael 1884, p, SJ,
Trombidioidea Banks 1894, p, 209; Southcort (957s,
p. 173; L982, p. 285; tinier card 1973,
p. 109; Welbourn 1983, p. #3; 1084, p, 135
Trombidia beider 1Y54b, pp. 539; 1979, p. 420,
Definition: Prostigmatic mites of generally owpid
or elongate form in post-lurval deutonymphal and
adult toctopod) stages. Wath one or Lwo puirs ol
dorsal propoedasomal sensillary setae in all mobile
sluges, eenerally in association with a crista or shield
if Octopod stages. bul erista and shield may be
rudimentary, vbsolete ar absent, Larva generally
rounded, heaapod, with ane or more dorsal
idiasomal shields, sensillary selac (one or two pairs)
borne by anteramost shield. Gnathosoma well
developed in all mobile staves, with mobile digits
as hinged blades, not retractile, not styliform,
Ocvtopod states cenerally wath genital acetahula
(suckers). Coxse | and Il, and Il and Ty,
contiguous on each side in octopod stages, | and
I] generally continuous in larvae, bul may be
Separate in larvae. Larvae heteromorphic to ovtopod
Stages. Larva with urstigma and anus. Octopod
stages predalory upon sinall arthropods. Larvae
venerally purasite upon invertebrates ond
vertebrates, Mires nevee fully aquatic.
Tyne genus Trenriidinm Fabricius, 1773.
Remarks: The larvae af the Trombellidae and
Johnstonianidae may have ove of more of the
following characters:
(1) vonae usually separated and urstigmea usually
separated from coxa L
(2) idtosomal setae usually arise from expanded
basal plates
(3) usually many sensory setae on the Jemura and
gcnua of the lees,
Two family zroups may be distingurmshed among
these Larvae, as. follows:
Dorsal shield well developed, with 6 or B sere, including
one pair of senyillary setae, well developed, hur never
clavare ov thickened, Caxae of legs separated, Urotlema
allached (o posterior margin of coxa 1. Supracoxalae
present ar absent. Palpat tibtal claws generally well+
developed: pibareare or wifaredte cays
: ‘Trombellidae lamily group
iaorsal shicld phisent with eight sete, bu; may be only
moderately sclerotized; a small separare anterior pam may
be present, Generally two pairs of scutal sensillary sctac
(one pair only, in On’ genus), of whith one pair may be
enlarged in veatrit) part, or-even clavate. Anterior pair ol
sensillary setae and their alveoh may be ridimentary
Coxae (and Tamay be joined of separate on each side,
Urstigsia usually free of coxa — may project laterally
between the coxae, or be altiched io rhe anerior bender
28 RV SOGUTHCOTI
at a separate coxa 1 Clerterhenenbived), Supracexalac
present or absent, Palpal bial elaws generally small,
bifureate or with a single claw, or may be teplaced by
vlonpate setae, not clawelike,
coe eee cece ey Pontily Sotrnstonmiaritag (ys yer
considered a single family, witht
several suibfamnilies, as fir as
knowledge of the larvae is con-
cerned),
As can be seen, there are many shared characters
between the two groups. Nevertheless, at the family
level (here is rarely dilficulty in placement.
Trombellidae family eroup
The Trombellidue (amily group here teludes the
Trombellidac, Chyzeriidae and Audyanidae, fam,
nov. These are separated as follows:
Key ra the larvae of the Trambellidae family vroup
|, AM setae lacking. Dorsal propodosomal scurum with
f setae. Bae segmental formula 7, 7, 7 Pedotgrsal
claws 3. 3, 3. Palpal tibial claw trifureate, Lateral
surface of cheliceral blade with myAity fine teeth,
Supracosalac present, Eyes 2 4 2 -
: | Chyzeriidae
AM seine present. Dorsal proradosorial sculum with
# sctac. Log segmental formula 6, 6, 4 or 7, & &
Pedaovarsa] claws 1, 1,1 or 1, 1,2 of 3, 2, 2 Palpal
tibial claw biturcate, |.areral surtace oF chelicerul blade
amooth, normal. Supracoxalae absent. Eyes 2 2
RT SSCN Ee res wlll BAe wou ten .
Leg seginenial formula 6, é, 6 Pedotarsal claws 1, 1,
tort, 1,2. None of se utal setae shart and clavale,
Coxalae antl palpfemoralae normal, setulose Eyes 2
: Trombelliday
lee segmental formula 1. 6, 6. Pedotarsal claws 2,
2, 2 Some of sental sorac short and clavate, also
coxala 11 a TH, sl palpinirrgraly hyes absent
.. And yanidae
bea
occce yo beee
Family Jrombetlidac
Partial synonymy
ireobelljpac Por 1935, p. LOS! Wamersley 1937,
p. 73; 1954u, p. 117; 1954b, p. 121. Vercammen-
Grandjean 1973, p, 109, Vercammen-Grandican
ev it, W974, ys. 245,
Trombelfidue Peder 1995, pp. 90,67; 1979, pp. 42),
422, Southcott 1982, 7, 289; 1986a, p, 145.
Thrombellinge (sic) André 1960, p. 315; Roba
1948, p. 453, (all ad p,).
Ihrombeltlidae Robaux 1973, p, 124,
Trombstoidew Peider 1979, pp. 421, 422 (ad p.)
(nom. mud.)
Redefinilion; Adult and deutonymph:
Lrombicioidea in which the prapodosoma. cither
lacks a cristu or fas oaly a rudimentary eristay one
pair of sensillary setae (trichobothria), Eyes 2 +
2, sessile, Luiosoina niay bear large plaques, which
may be arranged in columns on upper surtace of
whosoma. Idiosema nor attenuate or waisted,
Larva: Trombidinpidea with one doréal
propedosomal scutum, which projects anteriarly io
a narrowed extension or nasus. Dorsal scutum with
eight setuc, comprising 2 Al.s, 2 Pls, 2 AMs, aril
2 well separated sensillary setae, placed between
ADs and PLs. Eyes 2 - 2. Leg segmental formula
6, 4,6, CoNae Separated. Pedocosal formula 2, 1,
hor]. t, 1. Pedotarsal claws 1, 1, bor t, 4, 2.
Supracosalae absent.
Type genus Tromihella Berlese, [X87-
Remarks: Vhe 'trombellidac, as restricted, includes
all the genera listed curliyr (above) lor the Mamily,
except Chyzeria, Nothatrambicula, Audyana and
Ralphavdyna, The larvae of remaining genera muy
be separated as in the following key:
Key in larvae af Trombellidae
1, Pedotarsal claws i 1, 1. Claws sample. - itz
Pedotarsal claws J, 1, 2 (in 7 and If the single claw
is apically HrifDICATE) oes cess ees bteees renee
te
Chelicerae compact, the combined chelobases about
as long as wide, Sculal sensilla generally well behind
the level of AL sculale ....... Trombella Berlesc
Chelicerae elongate, the combined chelobases abour
twice as lone as wide, Scutal sensilla only a@ Jiihe
behind level of AL seulalae.. 0.22.0 ...2.-2,.--.
HE A tare. Nothratrombidium Womersley
3, Nasus of scntum sinall, largely accupied by the bases
of the AM scutatac, anid with & deep constriction
behind. Lee tibia Uf) with a large solenoidala -
. Wamerstepir Radford
asus al ,cuLuteh large, Ihdndular, its lateral borders
conbnuous with anterokuteral borders of scutum, with
at most only minor constriction. Leg Ghia Ui wirhout
A large solenmdala . 2.6.6 eee ee
Durenia Persamnmen-Gratidjean.
Nathratronbidiun Womersley, 1954
Redefinition of larva: Trombellidae, Sensillary
setae arise behind middle of scutumi, a litle behind
level of AL seutalae. Coxal setal formula I,
Pedotarsal claws lL. 4, lL. Chelicerae bases long and
Slencers combined elrelicerde bases whoul twice as
lung as wide. Palpi long and slender, palpal tibial
claw small, with two minute terminal nearly
apposed prongs,
Type species N. ofiaruet (Berl) Cadilil
Nothrarennibidinin treali sp. wy.
FIGS | A-E; 2 A-(; 3
“Larva of undetermined genus”, treat (1975, p, 236),
Holotype (in American Museum of Natunil
History) mounted in Hoyver's tector,
identificanion ACB760, somewhat damaged, found
dead “under right forewing of Spaeloris clandestina
\(Harcs)| 68-39 co |Lepidoptera. Noctuiditel
“Tynngham, Masstachusetts], U.S.A. 77 Sept, 196s,
A. E, Treat - (31; 29)", taken (O10 pm
MITE CT ASSIFICATION 24
PARLE 1. Meprie due nf pve species of larval Nothrotrambidium.
TNF AW PW SRK ASB PSB L Ww
SL ieutl ap. riy. 36 47 02 1s 78 44 f22 76
Holorype
Novrithorin (Beck) 90-109 fl
({rani Feider, 1958b)
AP AM AL PL AME SE DS MDS PbS
Nv treat! 45 16 16 29 14 98 33.62 38-49 48 53
Toh Tel Oel Til Tal | Trochlt Pell
No treat? 50 10D 33 3 1s] ad tov
N. oftierun 44 lle 5s 22 IAN 41 122
Gell TiU hat yp Trach Tell Gell
NL eaté 52 43 (25 o4 1s 2
No atiorunt 458 TOO 136 fl 145 47
Tilt Fabtt TH / Gel Tilt Gell THN Gell
N. trealt 14s Id {49 1,60 234
No oreo T&R 1ay (2.10) (0.72) (2.81)
©“ {N = distance from ablerior tip of scutum ta level of the AM setge tsee Southcol! 18966),
7 Omitting glaws and pedicle of the tarsi.
Description of holotype larva: Colour in life not
available; the specimen was found dead by Dr
A. E. Treat (see Treat 1975, p, 236), who advises
(pers. comm, 1983).that the dead mite was orange.
Length of idiosoma (partially engorged) 390 pm,,
width 255 jam; total length of animal from tip of
chelae bases to posterior pole of idinsoma 485 jam.
Dorsal scutum ovoid, narrower anteriorly, lateral
and posterior margins somewhat flattened, but
generally smoothly rounded.
Sculal scobalac as in definition; scural sensilla
behind middle of sewtutn, a file posterior to AL
sculalae, somewhat separated from each other;
scultalac tapering. short, witl slight setules,
Metri¢ dats are as in Table 1.
“yes 2 + 2, sessile, each lateral pair set on oval
plate, separated from dorsal scutum. and placed
between levels of AL. and PL scutalae in specimen,
Corneae oval, anterior 13 pm in longest diameter,
posterior 15 zm.
Dorsal wiosomial setae slender, tapering, slightly
blunted at tip, with minute selules; arising from
normal seta-bases (annul); arranged 6, 6, 6, 6, 5,
2, toral #1.
Venter of idiosomia with coxa I and IT moderately
separated on each side. Sternal area with a pair of
svobalae, between coxae Il, slender, tapering,
pointed, with small setules, 31 zm long. Between
coxge I] a further pais, similar, 33 pm long, but
with slightly more outstanding setules. Behind Icvel
of coxae U1] are 38 setae, slender, tapering, in
irregular transverse rows, 15-49 pm long,
lenethening posteriorad: {he more anterior of (hese
similar to the last-named pair, more posteriaracl
becoming smoother, similar to the posterior dorsal
idiosomalae. Anus (uroporus) 35 am long by 9 pm
wide, with two slender valves and a crumpled lip.
Coxalae 1, 1, 1. Coyala [ anses near AL angle
af coxa, slender, tapering, pointed, with faim
setules, 47 pm long, Coxala Il arises anterior and
lateral to central point of coxa, similar to {, 34 p,m
long. Coxala HI arises anterior to centre of coxa,
simular fo | and IT with adpressed setules, 32. pum
long.
Lez segmental formula 6, 6, 6. Legs lang anc
slender, femoral to ubial segments mote or less
cylindncal; tarsi clongate, spindle-shaped, Leg 1 510
pm tong, IL 445 gm, HW 595 pm (lengths inclade
coxae and claws). Pedocoxal supracoxala absent,
Tarsus } LST pm long by 20 pm high where thickest,
W125 pm 2 20 jun, Ub 146 jain. 18 um Clengths
exclude claw anid! pedicle). For other measurements,
see Table 1. Tarsi bear small, faleiform, stinple,
slender, single claws (in Specimen broken in R Tey
Il, missing in 1, leg 0),
Chaetotaxy of legs: leg scobalae (normal setae)
slender, tapering, pointed, with light setule
formation, Setac of leg scgments are indicated in
Fig. 3, A number of specialized setac present on leg
segments, in addition to scobalac, distal ts
trochunters. These include a number of spinalae
(eupathidalae), These and orher lee setae are
identified in Fig, 3 as far as possible, but since, in
the damaged specimen only three reasonably
cotiplete legs were avilable (L leg L, R legs HW and
11), it has not been possible always to identity them;
ina number al cases only the setae bases remain.
Vestigiogenualae present: VsGel.85pd (7 um long),
VsGell.84pd (6 pm), Vestigiotibiala: VsTil 86d (7
pin). Solenotarsalae aré present to | and 1]: sotal,1Wd
R. V. SOUTHCOTT
MITE CLASSIFICATION M
(37 pm), SefallJ8d (22 ym); fanvulus is present to
| and Uf: FaTal.36d (4 em), Falall.36d (4 om), (Tarsal
lengths nteasured jo origin of pedicle; for
explanation of coding symbols see Southeotr
I96la,b, 1963),
Cmathosama (damaged in speamen, part of one
chela missing) smiull, elongate, combined chelac
bases to origin of cheliveral divit (blades broken or
missing) 84 um long, x 47 pm across, clongate-
pyriform. Galeala present, pointed, smooth, 1 um
long. Anterior hypostomala apparently absent,
Palpal setal foraiula 0, 1, 1, 3, 8; palpal coxala
("capitular sera”) apparently absent. Palpal
supravoxala absent. Palpfemorala (lorsal in
position, short, pointed, with few setules, abour 14
um long (? broken), Palpgenuala similar, dorsal anil
distal, ca 10 jaw long, Palpal tibialae and tarsalae
as figured, Palpal Libial claw small, slender, with
only a slightieleh and owo minute ventrally directed
prongs.
Remarks: The larva was found “on a ventral axillary
Membrane of the right wing” of the moth, which
had been raken at light. However, the mite has only
one dorsal scutum, The idiosamna contains a mass
of fungal hyphae (Treat 1975), “The mite's leus were
whitish... the mounted mite showed a clump of
mold mycelia with fruiting bodies on one side of
the idiosoma, This was white in the fresh,
unmounted specimen” {Treat pers. comm, 1983).
{The pallor of the legs is consistent with partial
drying of the mite.)
The body of the mite contains fungal elements
consisting of hyeline septate hyphae with numerous
intercalary chlamydospures. No frutting bodies are
visible in the mounted specimen, Whether the
fungal infection occurred before or alter the death
of the mite cannot be determined, bul the lalierc is
considered more likely. The fungus is placed im the
Fungi Imperfecti (G. Kominski & D. Ellis pers,
oomm.),
Nothrotrombidium is he only trambellid Jarva
in Which the chelicerae are elongate; presumably this
character has some adaptive value, if the Jinding
ofaN. read larva ou a lepidopteran indicates some
advantage in burreawing through deep layers of
scales.
Taxonomic position of N. ireati
The two known spevies of Nalarol/rombidium
Turvae nay be separated as follows:
Ja Til¥Ge tf > 2 about 29 ventral setae behind
Conae iif, veer eee eee NM. alianenm (Berl)
Ib Tif/Gel < 2; about 38 | ventral setae behind coxae
0 - N. treat sp. now.
Fig. 1. Nothrotrombidium treat sp, nov,
Remarks: There appears no doubt that N. treads is
congeneny with the larva of NM. olerwn as
described by Feider (19586). The urstigma is well
shown, so thal jhere is no douln as to its
trombidioid affinities, consistent with its general
appearance. Coxa | is shown well separated from
voxa H, which is i agreement with the structure
of Trombella and Chyzeria.
A pale of setae on the idiosomal venrer, anrerior
to couse Tis shown by Feider (1958b, Fig. 3}, which
may be-an error of interpretation of some fold of
integument. ‘The figures of the gnathosoma (his Figs
3, §} Show a pair of hypostomalae (palpal voxalae,
or tritorosirals sensu Newell 1957, p. 403) level with
the medial angles of the femora, which L have tat
been able to identify in M. freati, presumably
because | have only the single damaged specimen
available Feider (1958b) illustrates barbed dorsal
and ventral palpal femoral setac, as Well as a barbed
dorsal palpal genual seta, allhough his text omits
mention of the ventral femoral seta,
Family Chyzeriidae
ms Trombellinae, Thromhellinac auctarum, +
sup. (ad pr).
Chyzeridae (sic) Krang 1978, pp, 278, 304,
Definition: Trombidioidea in which the
propadosoma of the adult and deutonymph either
lacks a crista or has only a rudimentary crista,
bearing a pair of sensillary setac, Eyes 2 ; 2, sessile
Dersum of idiosoma produced into a number of
long processes, Idiosoma not elongate or waisted,
Larva with one dorsal scutum, with 6 setae: 2
ALs, 2 PLs and 2 sensillary setae, sensilla well
separated, arising at about middle af scutum.
Scutum with or without anteromedian nasus. Eyes
2 + 2, sessile, Leg segmental formula 7, 7, 7. Coxae
separated. Coxal setal Formula 2, 1, §, Pedotarsal
claws 3, 3, 3, the neolateral claws with widened Lips.
Dorsal and ventral idiosomal setae may have
expanded seta bases. Supracoxalae present to
gnathosoma and leg 1. Lateral surface of cheticeral
blades with many fine teeth. Parasitic of
Onrthoptera.
Typé genus Ciyzeriy Caiestrim, 1297,
Remorks: The two genera new included to the
Chyzenidae may be separated asin the following
key:
Key ta the gener of the Chizeridae
Dorsal scutum withouc hasus, Palpal iarsalae may be
vatiousty moditied wih able) bu mot long and Feather-
like nat _ Chyzerva Canestrini
larva, Holotype A Dorstl view, lees omitted soand trochanters. pde
podocephalic canal. B Dorsal view of tip of rizbt palp. C-E tdiosomal setae. C posteriur dorsal idiosomala. 0
sentral idinsomal seta of first row behind coxae Hl, E pusterior ventral jdiosomala. (All [igures ro nearest scale.)
32 R. V. SOUTHCOTT
Fig, 2. Nethrotrombidium treati sp. nav, larva, Holotype, A Ventral view, legs omitted beyond {rochanters, to scale
on left. B Ventral view of right palp, to scale on right. C Further enlargement of tarsus of right palp, not to scale.
Fig. 3. Nothrotrombidium treati sp. nov., larva, Holotype. Legs T, I, TE. Codes: L of left side, R of right side. av
anteroventral. pd posterodorsal, Sc scobala (normal type seta), So solenoidata, Sp spinata (eupathidala), Vs vestigiala.
Symbol A means that the seta is shown doubly, in both aspects for the leg. (Al! figures to same scale.)
33
MITE CLASSIFICATION
uM R. V. SOUTHCOTT
Dorsal seutiim with a premnent narrow Masus extending
abrupuly frous anterior border. Some palpal tarsalae lone
und feaiter-like -...... Notieatrombicu/a Dumbleton
Audyanidae fam, nov,
Definition: Trombidionea in which the
deutonymphs (adults are as yet Unknown) lack a
crista, Larve dorsal propodosomal shield present,
with anterior wotch: (wo sensilla at posterior edve
of shield, well separated, Idiosomal setae with
annulus produced to form a high papilla bearing
a small scobala at its (ip, arranged in groups of 2-12
on small plates of the idiosoma, Similar selae on
propodosomal shield and legs, Eyes.absent, Palp
with strong tibial claw, clenidium present on palpal
tibia as two dorsal spines, Palpal tarsus. clavate,
modcrately enlarged. Genital valves each with single
large, oval actubulum.
Larva with single prodorsal seuturm, with & setae:
2 AMs, short, clavate, 2 similar ALs close to the
sensilla, 2 normal PLs, sensilla present behind
middle of sculum, sensillary selae pointed,
somewhat thickened. Nasus lacking to scurum, Eyes
absent. Leg segmental formula 7, 6, & Coxae
separated, Coxal setal formula 2, 1, |; coxalae I
and II] clavate, similar to AM and AL seuralae;
palpal fernorala similar, TVarsal claws 2, 2, 2.
Supracoxalae absent,
Content: Audyane Womersley, 1954a.
Johnstonianidae family group
Family Johnstonianidue
Partial synonyiiy
dohnstonianinac Thor 1935, p. 108: Womerstey 1937, p. 76;
Thor & Willmann 1947, p. 221; Foider 19554, p. 75.
Johnstonianidae Newell 1957, p, 396; 1960, fh, 156;
Feider 1959b, p, 540; 1979, (9, 420 tulso
Johnstonianoidea), Vercamiuien-Grandjean 1972, p. 227;
1973, p. 110; Robaux 1973, p. 12h; Vercammen-
Cirandyean ef al, 1974, p. 245,
Definition: Adults: Small or middle-sized
iremnibidioids with or without propodosomial dorsal
crista, Propodosama with one or two pairy of
specialized or relatively unspecialized sensory seiae,
which may be set in trichobothrial pits or be less
clearly defined. Eyes 2 + 2 or lacking; if present,
on short peduncles. Skin comparatively smooth,
without prominences or Jarge sclerotized areas,
Idiosomal setue simple, nude or, if setutase, nut
vlaboratcly so; they commonly originate on small
plates or raised papillae. Genital acetabula 3 + 3
or 2 + 2, Pedotarsal claws without empodium or
empodium-like brush. Palp generally long and thin,
with or without tibial claw, with few or no accessory
spines. Paragenital scleriles presen! ur absent,
Predatory, may be associated with semiaquatyy
envirgniments,
Larvae: Propodosomal dorsal scutum with Ww
pars Ol sensory setae and four normal setae, ar wath
two pairs of setae, one pair sensory and the other
pais normal, sensory setae may be rudimentary or
fully developed in trichobothrial pits. Leg segmental
formula 7, 7, 7 or 7, 7, 6 or 6, 6, 6, Coxal setal
formula 2, 2, 3-4 or 2, 1, 20¢ 2, f, bor Ld
Coxal setae setulose, not highly modified,
Pedolarsal claws 3, 3, 3 or 2, 2, 2. Eyes 2 + 2,
sessile, or absent, Supracoxalae presem or absent.
Free-living predators oF ectoparasitic on insects,
at times semiaquatic
Type genus Jodmstoniaml’ Goerge. 1909,
Remarks; Thor (1935) defined the subfamily
Johnstonianinae as (translation by author):
“Body (abdomen) cylindneal, with pointed, simple hairs.
Crista well developed, with two arcolac in the middle
(or one distal) and Four (-two pairs) of sensory setae.
Anterior ro the thorax is an elongare triangular
projection (nasus), Eyes on short peduncles
(kurzgestielt). Palpi almost without, or with few, spines,
Legs of middling lenerh,
Type: Johnstanianay C. Fo George 1909 (svn.
Diplothrombium Berlese, SHO = Roheultia Qudemuns,
917",
Other genera placed in this subfamily by Thor &
Wilhnann (1947) were; Centrotrombidium Kramer.
1896, Diplotironrbium Berlese, 1910 (they did ot
accept iis synonymy with Johnstuniana),
Myrmicotrombium Womersley, 1934, and
Hirstthrombinm Oudemans, 1947, the last-nanyed
with type species Diplorhrambium wustratiense
Hirst, 1928. They removed Notothrombium
Stork4n, 1934 to a separate, new subfamily,
Notothrombiinae,
Myrmicotrombium was shown by Southcort
(1957a) to belong to the Brythracidac.
All of the genera listed were based on the adults,
exeept for Rohaultia, which was considered to be
a larva of JoAnsloniana (although this has been
widely accepted, Cooreman (1949, p, 10) pointed
out there was no certainty Uhal Reohaultio
hiungulurm Oudemans, 191 was the larva ol
Rhynchaolophus errans Jahnstan, and in fact
Rohauitia was not established to be a larval
synonym of Jo#ristoriana wotil Keer recorded anc
described the rearing of larvae of Jodmsroniana
thaxiria Feider, 1955 (Feider 1955a, 1958a)). Robaux
(1970) described larvae of J errans obtained by
experimental rearing.
Further genera which have been placed in the
Johnstonianidae are (see Vercammen-Grandjean
1973): Polpdiscia Methlagl, 1928 (larva),
Parawenhdek ia Paoli, 1937 (larva),
Crossethraembine Wome4r>sley, (939 (adult),
MITE CLASSIFICATION 35
Lasséenia Newell, 1957 (larva, deutonymph, adult),
Morcandreella Feider, 1957 (adult, deutonymph),
Charadracarus Newell, 1960 fadult, larva),
Pteritdopus Newell & Vercammen-Grandjean, 1964
(larva), Paraplothromibium Robaux, 1968 (adult),
Parachyzeria tHirst, 1926 (adult) was placed in the
Johnstonianidae by Southcou (1986a), Two further
genera placed in the Jobostonianidae by
Vercammen-Grandjean (1973) were
Nathoatrombicula and Grossia (= Chyzeria). Both
ol these genera have been placed here in the
Chyzeriidac. Porawernhoekia is exclhided, as of
doubtful affinity (see Southcott J96ta),
There remains a toral of mine genera in the
Johnstonianidae, known as Jarvae, fur which the
following subfamily groupings are proposed:
Johnstonianinac: Jufinstoniana f= Rohauitiad,
Diplothrombium, Centrotrombidium;
Tetrathrombiinae; Jetrathroabiunr, Lasseniinae:
Lassenia; Polydisciinae: Polvediscie, Preridopodinae:
Preridopus; Ralphaudyninac: Relpheudyne:
Charadracarinae: Charadracarus,
The Following is a key to the subfamilies of the
Johnstonianidae (larvae),
Key (o the larvae in the subfamilies of
Jobastonianidae
1. Yarsal claws 2, 2, 2, Anal selerites absent. Supracaxalae
absent to gnafthusoma and leew T........ 2.2262, 2
Tarsal claws 3, 3, 3. Anal sclerites present... 3
2. Leg segmetital formula 7, 7. 7 or 6, 6, 6, Posterior
sensillury setae of pradorsal scutuin without thickened
central part to shall, filiform (except in
Centrotrambidium where whe vensitlary setae are
terminally clavate), Sterial setae: usually a pair of
sclae between coxae ILL. Terminal seta of palpal rarsus
not cupathiditorm. Urscigma between coxac | and 11
not projecting away laterally (ram idiosoma, Coxal
formula 2, 1,1, Byes 2. + 2. Tracheae absent. Anterior
hyposiomala absent ...-_, .Johnstonianingae Thor
Leg segmental formula 7, 7, 6. Posterior sensillar
selae of prodorsal scurum with thickened part in
middle. Sternal setae numerous in area belween conde
Il and WI. Urstigma in a chitinous extension
projecting well lateral from idiosoma. Coxal formula
2, I, 2. Eyes absent. Tracheae present. Amerior
hypostomala absent. Terminal setae of palpal tarsus
nol eupathidiform ,,, ...Charadracarinae Newell
3, Lev segmental foroiula 6, 6,6. Coxul setal formula
2, 1, 2or2, 2, 3-4. Byes 2 + 2. Anterior sensillary
setae borne on a sniall selerite which may he separated
fram the main body of prodorsal scutum. Sternal
setae absent. A well-developed pore of @ “lasseaus”
ar “fassenio organ" present upon a small separate
sclorile auterint (6 coxa IL]. Suptacexalae present.
Lasseniinue Newell
Leg segmental formula 7,7, 7. Byes 2 + 2.....4
4, Attterios scutal sensilla in anrerior third of prodarsal
sculum, poskenor scutal sensiila in postertur third.
Stermal seme absent, Coxal setal formula 2, 1. 1.
Tetrathrombiinae subfam. nov.
penne
Anterior and posterior pairs of scutal sensilla both
in amerior (woshirds of prodorsal scutum -.. -- 5
5, Coxal setal formula 2, 1. 2. Sternal setae lacking.
Neolateral claws of pedoiarsi divided. Gnathoxomal
supracaxalae present... 2... 2. ee ee ee
--- oes... Polydisciinae Vercammen-Grandjean
Comal setal formula 2, 1, 1. Two sternal setae present
between coxae LIT, Neoluleral claws of pedotarsi
broadetied bul undivided. Supracoxalae present 6
6 Anterior sensilla of prodorsal soutum posterior to level
of AL setac. Bath pairs of scutal sensilla well
developed. Scutum with small nasus. Coxalae normal,
setulose. Tarsus HP extremely elongate and carrying
# dorsal row of Jong, feathered setae ..0.2,. 22. -
We. ey Lr .Pteridopodinae subfam. nav.
Anterior sensilla of prodorsa] scutumn anterior to level
of AL setae Anterior sensilla and seta¢ significantly
smaller than posterior, Scutum with large nasus. Conus!
setae (ubereulate with « subierminal bristle. “tarsus
Remark All subfamilies are known from only one
genus, except Johnstonianinae.
Sublamily Johnstonianinae Thor
Johnstonianinae auct. tad p, v. sup.)
Definition (larvaes: Prodorsal scutum with cight or
four setae; if with eight then these are 1wo pairs of
cach of sensillary se1ae and non-sensillary setae, if
with lour then with one pair of sensillary and one
pair of nonsensillary setae. If with Four setae hen
sensillary Selae clavate; if with eight setae then
sensillary setae filiform, and anterior palr may be
reduced. byes 2 + 2, each lateral pair on a small
ocular plate, which may be elevated into a tubercles
in Cennirarmnhidium postenor cornea may be
obsolete or lacking. Usually twa sternal s¢tac
berween coxac LL. Anal sclerites absent or weak:
if the latter, they are non-etiferous, Coxalae 2. |,
1; medial coxala I generally on a small pars medialis
coxae, which is rarely separate! from vesz.
Urstigma between coxae | and Il, not projecting
away laterally from cowae. Pedotarsal claws 2, 2, 2.
Galeala present, anterior hypostomala present;
reduced or absent, postenor hypostomala present,
Palpal tibial claw bifurcarc, Terminal seta of palpal
farsus not cupathidiform. Trachese absent,
Supracoxulae absent.
Type genus Johisionlana George, 1909,
Remarks: The following is a key jo rhe venerp of
the larval Johnstonianinae;
Kep to genera of larval Johnstameniiae
I. Scutum with four setae, posterior pair clavate
sensillary setae Vs absentalltegs.... ..- i
Se, .. Centrorrambidiamn Kramer, 1895
Seutum With cieht sete (lwo pairs sensillary, two pxins
fon-sensillaty, senoillary selae not clavate -.....2
% R, ¥, SOUTHCOTT
2. Anterior pair of scutal sensillary setae ar least one
third as long as the posterior pain VsGel II present,
VTL, HW, WE absent, Anterior wall of palpal trochanter
not fenestrated , ...- - ge tee Seat rhictie «
heat Johnstoniana George | Rakaultia Ouds.)
Anterior pair of scutal sensillary setae reduced, at
most One sixth as long as posterior pair. Vs absent
allsvaments Antenor wall of palpal trachanter may
be fenestrated ..., Diplathrornbserr Rerlese, 1910,
The key above has been drawn up principally
from descriptions of larvae allotted (o (hose genera
as Follows: Jofmistosiana from J dutiscuta Newell,
1957, £ waxiina Feider, 1955 (described by Feider
1958a), J. Rarghifensis Feider, 1958c, < veniripilesa
Feider, (958, J. errans (described by Robaux 1970)
Diplothrombium monoense and D. cascadense af
Newell (1957), D. molduvicum Feider, 1959a, D,
newelll Robaux, 1977; Centrotrombidium trom C
distans of Newell (1957), C. rameniense of
Vercammen-Grandjean & Feider, 1973, C
dichotomocoxala Vercaminen-Grandjean A
Cochrane, 1974.
‘Tetrathrombiinac subfam. nov.
Definition (larvae): Anterior pair of sensilla in
anterio: third of prodorsal scurum, posterior pair
in posterior third, Sensilla well-developed, sensillary
setae filiform. Eyes 2 + 2. Sternalae absent. Coxae
separated. Urstigma attached to anterior border of
coxa LI. Leg segmental formula 7, 7, 7. Coxal setal
formula 2, 1, 1. Pedotarsal claws 3, 3, 3 (lateral claws
may be reduced). Galeala, anterior hypostomala
and posterior hypostomala present, Palpal tibial
claw bifid.
Type genus Tetrathrambiumt Feider, 1955.
Remorks: Jetreihrombian is known trom (wo
species, 7. zuchvaiKini Feider, 1954b (type species)
and T. macronychus Yeider & Suciu, 1956, from
Europe. 1. zachvatkini was obtained as ectoparasitic
upon a plecopteran, and T muacronyehus ftom
lipulid Diptera. The adults of the genus are
unknown,
Lasseniinae Newell
Lasseniinae Newell 1957, p, 447; Vercammen-
Grandjean 1973, p, 110,
Lassenidae (sic) Vercammen-Grandjean
p, 236,
1972,
Definitian (larvae): Anterior scutal sensillary setae
bornc on a small sclerite which may be scparate
from main body of <enjum, Eyes 2 + 2. Sternal
selae absent, Anul sclerites present, setifernus,
Lassenus or “Zussenia organ” present (a small pore
on a small separate sclente anterior tO coxa JM).
Leg segmental dormula 6, 6, 6, Coxal setal formula
2, 1, 2 or 2, 2, 3-4, Coxalae setullose, (inmodified,
VsGel, Fl present, VsTil present, VsTill absent,
Pedotarsal elaws 3, 3, 3, Galeala, anlerion
hypostomala and posterior hypostomala preserr
Palpal tibial claw unidentate a: bidentate.
Stupracoxalae present.
Type genus Lassenia Newell, 1957
Remarks: At present Lasseniinae should be
restricted to one genus, Lassenia, known as larvae,
deutonymphs and adults. Two species are known
as larvae, both from North America. In neither case
was there experimental correlation between larvae
and aclepod stages, and correlation Was based
purely on strong presumptive tield evidence (Newell
1957). The larvae are parasitic on Diptera living in
subaquatic environments.
Vercammen-Grandjean (1973) included in the
“Lussenidae” Lassenia, Polydiscia Methlagl (known
only as Jarva), and Crossothrombium Womersley,
1939 (known only as adult). The status of whe last-
named (which has 2 + 2 eyes, contrary to
Womersley’s description) will be considered In
another paper, Polydiscia was made the type genus
of Polydiscinae (sic) Vercammen-Grandjean by its
author (1972 this subfamiilial status will be retained
here (se¢ below),
Polvdisciinac Vercammen-Grandjean
Polydiscinae (sic) Vercammen-Grandjean 1972, p, 236,
Definition (larvae): Dorsal propodgsgmal scwlum
well developed, with two pairs of trichebothrial
setae and four non-sensillary setae. Anterior pair
of sensillary selae originate anterior to AL. selae,
behind slight convexity of anterior border of
scutum: ho defined nasus preset; scutum markedly
waisted. Eyes 2 + 2, Sternul selae lacking, Anal
sclerites present, setiferous, Coxa — and Tl
contiguous on cach side, with urstigma berween
them. Leg segmental formula 7, 7, 7. Casal setal
formula 2, 1, 2; coxalae setulose, unmodified,
Pedotarsal claws 3, 3, 3; cmpodium thin, falciforim,
neolateral claws divided, Dorsul eupathidala |
companala present to tarsus | and II. Palpal tibial
claw greatly modified, reduced to seta without huok
structure, Galeala present, anterior hypostomala
present, posterior hypostomala absent.
Gnathosomal supracoxalae present.
Type genus Polydisciy Methlagl, 1928
Remarks: Known only for the species /? seuurnate
Methlagl, 1928,
Vercammen-Grandjean (1972) gave a careful
redescnption of Pobwdiscia squamata Methlagl, ard
MITE CLASSIFICATION IT
TABLE 2. Pedal seehaler formula for titeé species of larval joknstonianid and one kivdryphaatid larval, mites.
Lassenia Pulydiscia
lasseni! syudinatal
Lez [ u ul i il
Fe 10 10 10 6 7
Ge 8 8 8 4 4
Ti 16 17 15 9 9
Ta 50 43 40 21 21
Sub-rotals 84 78 73 40 4)
Totals 235 120
Plersigia Charadracares
limophila! delitescens*
il [ IL tu J u Iu
6 7 7 6 6 8 6
4 4 4 4 s 4 4
9 g i) 9 § $ 1
20 1k 17 7 22 17 12
3¥ 38 37 36 38 44 32
Mt 104
\From Vercammen-Grandjean (1972), re-atranged-
“Derived from the iWustrations of Newell (1960).
founded Polydisclinae on the sole genus Polydiscia
on the grounds of:
(LY limited size of palpotibial claw
(2) presence of subterminala {+ dorsal cupathidala)
and parasubterminala {= companala to dorsal
eupathidala) on tarsus If
(3) “tabulation of leg setae is very much like that
of certain water mites . . "and commented “Thase
vharacters seem to imply a high ancestry to this
Benus, 4% does the peculiar shape and
ornamentation of the scutum, which interestingly
links the Lasseninae (sic) lo the Hydryphantidae”
(Vercammen-Grandjean 1972, p, 234),
In 1973 he listed (without discussion) Polvdiseia
in the Lasseniinae (possibly due to a publication
delay),
However, the grounds advaneed by Vercammen-
Grandjean for the separation of the Polydisciinae
are worthy of discussion, es they arc relevant wo
subfamilial classification within the
Johnstonianidae. Criterton (1} is true, but by itself
does not appear to be a justification for subfamilial
status, Criterion (2) also applies in ee. Lassenia
fasseni Newell (see Newell 1957) and £ seuterlata
Newell (see Newell 1957).
Criterion (3) is of more interest, and is an
expression of af increasing tendency among
specialists of the prostigmatic mites to use the
numbers of normal setae (scobalae) on the lee
segments in higher classifications. These numbers
imay be expressed in a “pedal seohalar formula”,
as his been done eg, by Vercammen-Grandjean
41972}, Robaux (1977), This formula, for four
species of trombidiform mites, ix shown in Table 2,
From Table 2 it can be seen that there is a decrease
in pilosiy through the four genera Lassenta,
Pelydiscia, Piersigi# and Charadracarus, It
reduction of the number of leg scobalae is
considered to be derived then Lassenia is the most
primitive of the four genera listed, and
Charadracurus the most derived, These remarks
apply only to absolute numbers of scobalae, In the
case of the genua, however, Polyeiscie has fewer
setae thin Cheragracerus ac least for geng |,
On the other hand, if one considers the numbers
of idiosomal scobalae, of the larvae, the relationship
ig reversed between Lassenia and Charadracarus.
Thus Charadracarus larvac have a large number of
setae in the intercoxal area betweety coxae IL and
II, while Lassenia larvae have none,
It would appear, therefore, that any conclusions
abour the phylogeny of the various johnstonianid
subfamilies at Icast, based on the degree of pilosity
of the larvae, should be treated with caution. Other
characters must be introduced into such an analysis,
Pleridopodinae subfam_ nov.
Plecidopiidae Feider, 1979, pp. 420, 421 (wom, wuely
proposed without definition ot key).
Definition (larvae) Prodorsal scutum well-
developed, with anterior nasus and eight setae; two
pairs of well-developed sensilla, anterior pair
originating behind level of AL scutalae, Sensillary
setae setulose, not expanded, may be plumose, Eyes
2.1 2, Sternal setacc two, between coxae It. Anal
sclerites present. Lassenus present, Coxa¢ | and If
contiguous or nearly so on each side, urstigma well
developed, Leg segmental formula 7, 7, 7, Coxal
setal Forniula 2, J, 1; coxalae normal, sctulose.
VsGel. IE, Vail present; VsTill absent. Tarsal claws
3, 3, 3) neclateral claws with distal widening, Tarsus
IL elongated, carrying dorsally a row of plumose
s@laej similar setae on tibla IL. Cheliceral blades
with many fine teeth and tubercles. Palpal
trochanter not fenestrated; palpal tibial claw
strongly bifid. Supracoxalae present.
Type genus Pleridopus Newell & VYercanimen-
Grandjean, ISf4
Remarks; The Pteridopodinae at present contalis
only the genus Preridopus, known for two species
of larvae (adults are nol known) from Atnica: P
auditor Newell & Vercammen-Grandjean, 164
(type species) atid P pseudhaennemunia Newell &
Vercummen-Grandjean, 1964.
Ralphaudyninae subfaim, poy,
Definitien fierveae)s Prodorsal scutum well-
developed, with broad nasus; erght setae, Cour
3% R, ¥, SOUTHCOTT
sensory, Anterior pair of sensilla anterior to level
of Al. scutalae, smaller than posterior pair
Sensillary setae wot enlanyed. Eyes.2 1. 2. Sternal
setae! two, hetween coxae Hl Anal sclerites present,
setiferous Coxae land [I contiguous on each side,
With (he urstigma set laterally between them, Lee
segmental formula 7, 7. 7. Coxal setal formula 2,
1, 1: all coxalae modified, tubercular, with
subterminal bristle in only known species. Tarsus
[IT only moderately elongated, does not carry a row
ol plumose setae alang dorsum, Tarsal claws 3, 3,
3; all claws widened, but undivided. Galeala,
antenor and posterior hypostomalae present. Palpal
tibial claw bifid. Supracoxalae present.
Type genus Ra/phavdyna Vercammen-Grandjean et
al, 1974.
Remarks: The Ralphaudyninae contains only che
genus Rulphaudyna, known onty for ils type species.
Ralphaudyna Vercammen-Grandjean ef al, 1974
Definition: With the characters of the subfamily.
Ralphaudyna umamiensis Vercammen-Grandjean
ef al, 1974
Remarks; Through the kindness of Mr W. C.
Weibourn, Acaroloegy Laboratory, Slate University
of Ghia, | have examined three specimens which
conform to this species. Collection details are as
follows:
Japan, Shikoku, Ishizuchi National Park, Omogo
Uly (sic), 700 m, IS-25.Ni7I9BO, ex Tachyetnes
mbustus [Anders Rhaphidophoridas, Grylacri-
doidea] S, & J. Reck, codes WCW 81406-9, -2,,-9
local identifications (RVS} ACB727A, BC.
Significant morphological features have been
discussed on p. 26, leading to the above taxannmic
placenent,
Metric data of scutum and legs of these three
species ave provided jn ‘lable 3.
The dorsal scutum is shown in Fig, 4C, The
anterior sensilla have the rypical “half-lidded"
appearance of (roinbidiod scutel sensilla, The
stetrialae are between coxae IIE, Jong-conical with
faint adpressed setules, GO zim long. The palpal ribial
claws are Well-cleti: the tines are separated, and in
the qgirrect orientijion can be seen te be angled
about half-way along their length (Fig. 4A, B).
Remurks on biology: The finding of larvac of this
species as ectoparasites on a gryllacridoid (cave
cricket) is of considerable interest. Many of the
larvae of the Johpstonianidae have been taken as
ecloparusiles upon water associated insects, eg.
upon tipulids, or even upon rhe aquatic pupae of
wait beetles uf an unnamed family (Newell 1957),
There i a superficial resemblance between
Ralohaveyae tarvac and Chyzeria in the highly
modified ooxulae ul at least same species of
Chyzeria larvae (see Southcott 1982),
Charadracarinue Newell
Charauracarinae Newell 1960, op.
Grandjean 1973, p. Ud.
Definition (larvae); Amerior propodosomal dorsal
scutum well-developed, with a sharp nasus and
slight-cvidenee of acrista; with eight setae: two pairs
of sensillary setac, two pairs non-sensillary. Anterior
pair of sensillary setae little different from sculalae,
without a well-developed typical alveolar pit,
Posterior sensillary setae with expanded middle
parc, arising from approximately norniul
trichobathrial pics. Ocular sclerites and eyes absent.
Sternal setae numerous on yventer of idiosoma in
area between coxve It and 1, Anal sclerites absent,
Coxae Land I] contiguous on cach side. Urstigma
well-developed, in chitinuus extension on lateral side
of coxa bt. Ley segmental formula 7, 7, 6. Coxalac
2, I, 2; coxalae normal, tapering, selolose.
Pedolarsal claws 2, 2, 2, Galeala presem, anterior
hypostomala absent, posterior hypostomala present.
Anterior wall of palpal trochanter not fenestratec
Terninal setae of palpal tarsus not cupathidiforn.
Supracoxalae absent.
Type genus Charadracarus Newell, (980,
157; Vercaummen
Remarks: Chavadracarinae includes only the genus
Charadracarus, with two species in North America,
C. hurdi Newell, 1960 and GC. delitescens Newell,
1960, and two. European species, GC grandjeani
(André, 1930) and C. aelleni (Cooreman, 1954). The
larva is known only for C. delitescens; correlation
between (he larva ancl adult for this species appears
to have bee based on strong evidence of
association in the field, tagether with the
morphological similarities of the adults and larva,
without evidence of experimental rearing. ‘There is
no present evidence to dispute the proposed
correlation; the larva is clearly a member of the
Johnstonianidac on other grounds.
Acknowledgments
Lam indebted lo Dr Asher E. Treat, U1S.A,, for
sending the specimen of Nothrotromibidiunt treats,
and for additiinal collecting information; and te
Mr W. C. Welbourn, Acarology Laboratory, State
University of Ohio, for the specimens of
Ralphaudyna amamiensis, | thank Mrs Geraldine
Kominski and Dr David Fllis, Mycologisrs,
Adelaide Children’s Hospital, for identification of
the fungus associated with MN. treats.
39
MITE CLASSIFICATION
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A Gnathosoma, dorsal. B Gnathosoma, ventral, C Dorsal scutum. D Part of coxa LIL and the two sternalac. (All
to scale shown.)
References
ANpRE, M. (1930) Sur unc nouvelle espéce francaise BeRLFSsn, A. (1887) Trombella glandulosa Berl. n. sp. (et
dacarien, appartenant au genre Typhlothrombium genus) (with plate). Acari, Myriapoda et
Berlese. Bull. Mus. Nat. Hist. Nat, Paris (2) 2(5), Pseudoscorpiones hucusque in Italia reperta. Number
527-53). 40, Pr. 2 (Vol. [V), (A. Berlese, Patavii).
(1958) Acariens thrombidions (adultes) de (1888) Acari Austro-Americani quos collegit
Angola. Publ. cult. Diamang No. 35, 1-125. Aloysius Balzan. Boll. Soc, Ent. Ital. 20, 171-186.
(1960) Contribution a l'étude des thrombidions (1902) Descrizione e figura della Trombella
Indochine. Acarologia 2(3), 315-326, atiorum n. sp, Riv pat. veg. (1) 9, 127-128.
Banks, N, (1894) Some new American Acarina. Trams, (1910) Brevi diagnosi di generi ¢ specie nuovi di
Amer. Ent. Soc, 21, 209-222. Acari. Redia 6, 346-388,
MITE CLASSIFICATION 4l
HesHeiiy da & Kier, J. AQ (1942) Veinticinoa especies
tuevas de Trambidiideas en Colombia, &ivy, Acad.
Culomb Cienn Exact, Fis-Ouim. Nat. S(LT), 110-127.
CANES iim, G. (897) Nuovi avaraidei della N. Guinea.
fermeészerm, biz, 1, 461-474.
LewOREMAN, |. (1949) Note sur dadiastenand ercons
(lohuston)(Acarien, Trombiditdae), Bull, Inge A. Sey,
Nal. Belx. 2542), 1-16
— (1934) Notes sur quelques acariens de la faune
cavernicole, frd, 30(34), 1-19.
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42 R. V. SOUTHCOTT
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(1939) Further noles on the Australian
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(1954a) New genera and species, apparently of
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S, Aust, Mus. 114(2), 121-128.
A NEW LARVAL MITE (ACARINA: TROMBIDIOIDEA)
ECTOPARASITIC ON AN AUSTRALIAN CENTIPEDE,
AND THE TROMBIDITDAE RECLASSIFIED
BY R. V. SOUTHCOTT*
Summary
Wondeclia centipedae gen. novy., sp. nov. is described, as an ectoparasite on the centipede Rhysida
nuda (Newport) (family Scolopendridae) from north Queensland. It is the first larval trombidioid
mite known as a centipede ectoparasite. The mite lacks eyes, and is unusual in the posterior
displacement of the anteromedian scutal setae and their wide separation, the anterior displacement
of the posterolateral scutal setae, the thickening of the anteromedian, anterolateral and sensillary
scutal setae, and the excavation of the anterior scutum's anterior border.
A new subfamily, Wondecliinae, is erected for the genus, which is included with the Trombidiinae
and Allothrombiinae in a restricted family Trombidiidae.
Metric characters for shield and leg characters customarily used in describing and specifying larval
trombidioid mites are analysed by correlation methods. A moderate degree of positive correlation,
above twice that of random expectation, is present throughout the sets of comparisons of the groups
of variates utilized. There is no excess of negative correlations.
KEY WORDS: Taxonomy, Wondeclia, Wondecliinae, centipede, Queensland, Acarina,
Trombidiidae, Trombidioidea, correlation.
A NEW LARVAL MITE (ACARINA: TROMBIDIOIDEA) ECTOPARASITIC ON
AN AUSTRALIAN CENTIPEDE, AND THE TROMBIDIIDAE RECLASSIFIED
by R. V SoutHcoTT*
Summary
SoutHcort, R. ¥. (1987) A new larval mite (Acarina; Trombidividea) ectoparasitic un an Australian
cenlipede, and the Trombidiidae reclassified, Trams. R. Suc. S. Aust. M11()), 43-52, 29 May, 1987.
Wondeclia centipedae gen. n0V., sp. nov. is described, as an cetoparasire on the conlipede RALSside
nuda (Newport) (family Scolopendridae) from north Qucensland. Ii is the first larval irombidioid mite
known as a centipede cctoparasite. The mite lacks eyes, and is unusual in the posterior displacement of
the anteromedian scutal setae and their wide separation, the anterior displacement of the posterolateral
scutal setae, the thickening of the antcromedian, anterolateral and sensillary scutal setac, and the excavation
of the anterior seutum’'s anterior border.
A new subfamily, Wondecliinae, is erected for the genus, which is included with the Trombidiinge and
Allothrombiinae in a restricted family Trombidiidae.
Metric charactérs for shield and Icy characters customarily. used in describing and specifying larval
trombidioid mites are analysed by correlation methads. A moderate degree of positive correlation, above
twice that of random expectation, is present throughout the sets of comparisons of the groups of variates
utilized. There is no excess of negative correlations..
Key Worps: Taxonomy, Wandectia, Wondecliinac, centipede, Queensland, Acarina.. Trombidiidae,
Trombidividea, correlation.
Introduction
Centipedées and millipedes are well known hosts
for commensal or phoretic mites of a number of
families (eg. Berlese 1882, 1910a,,b; Vitzthum 1941;
Evans 1955; Domrow 1956; Rack 1979), André
(1943) considered that some of these relationships
verged on parasitism, a suggestion confirmed by
Shiba's (1976) record of a larval trombidioid mite,
Milliotrombidium milliopodum Shiba, ectoparasitic
on millipedes in Malaya. However, tarval
trombidioid mites seem not to have been found on
centipedes (see eg. Oudemans 1912; Thor &
Willmann 1947; CloudsleyThompson 1968).
A new genus and species of larval trombidioid
mite ectoparasitic upon a north Queensland
centipede is described below and its taxonomic
position within the superfamily Trombidioidea
discussed. It is placed in a new subfamily of a
restricted family Trombidiidac, and some comiments
are made on its biology.
A key for the separation of the subfamilies of the
Trombidiidac is presented.
Seta terminology follows Southcort (1961a, b,
1963, 1986a, 1987),
TAXONOMIC DECISIONS AND ACCOUNTS
Superfamily Troimbidioidea Leach
frombidides Leach, 1815 (restricted)
Trombidioidea Leach; Southeort, 1987.
Definition: As in Southcort (1987).
“2 Taylors Road, Mitcham, S. Aust. 5062.
Family Trombidiidae Leach s, str.
Trombidides Teach, t814 (restricred)
Trombidiidac Sourheort 1086c.
Definition: As. in Southcott (1986e).
Type genus /hombidium Fabricius, 1775.
Key to larvae of subfamilies of Trombidiinae
1, Anterior dorsal sculumn more of less rounded anteriorly,
without anteromedian excavation, AM setae of anterice
dorsal scutum unmodified, arising towards anterior
pole af seutum and in front of all other setae of
seultini, Sensillary setae slender, tapering, aor clavate,
AM setae displaced posteriorad nearly to mid-level of
shield, short and clavate watlr long selules. AL setae
The anicrotnost setae of scutum, arising near AT, angles.
Sensillary setac clavate with strong setules, arising
hehind level of PL setae, yes absent... 2, -...,
Ny .... ... Wondecliinae subfam. now.
2 Urstigrs Oval® oe ees Trembidiinac
Urstigma circular? .........20.- Allothrombiinae
Remarks; With the discovery and formal
description of Wondeclia centipedae and its
placement in the subfamily Wondeclliinue, phe
subfamilies Trombidiinae and Allothrombiinac
Thor, 1935 {see the discussion in Southcott 1986c),
together with the Wondecllinae, should be plaved
ina restricted lamily Trombidiidae, as redefined
*This distinction is teutative. See the discussion in
Southcor ¢1986c).
At R, Vv. SOUTHCOTT
(Southeatt [986c), That definition applies to all
known adulls and deutonymphs of the
Trombidiinae and Allothrombiinae, but may need
inodification lor the Wondecllinae, when the pasr-
larval forms became known,
Subfamily Wondecliinse subfam, nov.
Definilion of tarvee: Eyes absent. Anterior dorsal
scrim without nasus:; with anteromedian
excavatiom AL seural setue ul antensdatent! angle
ef scutum; AM setae well separated, displaced
posteriorly nearly to level of middle af scutunt; PI.
setae displaced anteriorly to level of middle of
sculaim, Sensillary setae widely separated, clavate,
selulose, All coxal'setae setulose. Tarsal claws 3, 3, 3.
Type genus Wondeclia gen, nov.
Genus Wendectia pen. nov.
DeAnirion of farya: Wondechinae. Anterior
prodorsal shield approximately square, posterior
prodorsal scuijum # transverse oblong. Scutal
sensilla at about Jevel of junetion of third and
fourth quarters of sentum. Posterior dorsal seutum
with four normal, setulose setae) Tarsal claws
falciformn, simple, Gnathosoma with compact chela-
bases, Cheliceral blade with 2-4 barbed teeth.
Guleala robust, sinuous, expanded near is base,
unbranched, Palpal tibial claw bifid. Two large,
setnlose, posterior hypostomalae (tritoristral setae).
Type species Hordecha Centipede sp. nav.
Wandeclia centipedze sp. nov.
FIGS. 1-6
Deserplion ul larva Wrom Holotype SAM = N19879,
supplemented by other specimens). Yellow in life.
Idiosoma ovoid (Vip. 1), somewhat Mattened dorso-
venttally: length (mounted) 710 «am, by $70 pm wide
Anterior dorsal scupum widening a little
posteriorad, with (runcate anterolateral angles and
rounded posivrolaleral angles (Fig, 3Ad; all sides
somewhal concave, with anterior excavarion the
deepest, but owing to anterior edge being benr
somewhal forward and down, apparent depth
of uni¢rioe excavdtion may appear slight.
Anteromedian scutal setae short, asymmetrical,
with about four long, strong, pointed setules: (hese
setae nearer co edges of seutum than to. centre.
Anteralateral scutal setae tapering, poiuted, with
long, strong, pointed setules. Posterolateral scural
setae similar, but more slender, near middle of
lateral burder, causing o& slight conveaity there,
Sensillary setae mosr posterior setae af shield, long,
with long, strong, pointed secules, orifices facing
postcrolaterally (Fig. 3A), Ridge of chitin runs trom
AL anele of shield ta the annulus of AM seta,
Shield porose.
Posterior dorsal scutum: porose, anterior border
with a slight anterior projection, lateral borders
convex, proyceling anterolaterally, posterior bordel
straight or sinuaus; medial two setae anterior to
posterior pair, and nearer to middle of scutum,
posterior [Wo setae nearer fo lateral borders, All
selar with auislanding setules,
For metric details of seuta and legs, see Table L.
(See Fig, 4 for interpretation of the setae of the
scuta, and the conventions of seta and other
cadings,)
Dorsal idiosomal setae are pointed, with strong,
barbed setules, and arranged in rows of 2 (between
leve} of the two prodorsal shields), 4, 6, 10, 6, 2,
2; otal 32 (Fig.. 1). Veritral surface of idiosoma lacks
setae between coxae | and [1]; a pair of setae between
coxae TIL, slender, potmed, barbed, 37 zm long,
Behind level of coxae 111 about 27 setae in irregular
transverse rows, similar to preceding, becaming
longer posteriorad, 20-42 xm long.
Legs short, moderately robust; lengths (including
coxae and claws) 1 260 pm, 11 230 pm, U1 240 pm.
All coxalae long, pointed, strongly setulose.
Lateral cuxala 146 ym long, medial coxala | 37 arn,
anterolateral coxila 45am, posterolateral coxala
Il 39nim, coxala tll 27 wim, Urstigina large,
prominent, approximately circular, external
diameters L2 ~m by 10 xm. Lee scobalac similar to
coxalac: Irochanteral scobalar formula 1, 1, 1.
femoral 5-6, 4, 4, zenual 4, 2, 2, tibial 6, 8, 5. Pedal
supracoxalae absent.
Specialized setae of legs (excep iarsi} as follows:
SoGel.20pd (16 wm}, SoGel.40d (17 am), VsGel 7d
(4 um), Sofil.2?pd (134m), VsTUS9d (4 jem},
SoTL. 69pd (2 pm), SoGell.2dpd (29 zm,
Sofill3ipd (2 pm), SoTi.67pd (10 im),
SoGell1.46d (18 pm) (Rigs 5-6).
“Tarsus | and Il each with a large ceniral dorsal
solenvidala, Specialized setue eodings SoTal.44d
(4 pm), FaTal.49d (3 pm), Solall.44d (1 pm),
SoFall.33pd (4 yan), Peetarsal fornia 1, 0, 0, Tarsal
claws nermul, robust, subequal.
Gnathosomy robust, compact, each chela base
from above irregularly ovoid, length §7 pm from
lips of extruded chela digits to pasteriar pole of
bases; combined chela bases 5$ am wide. Chela
digits (blades) sharp-pointed, with 2-4 (usually
retrorse dorsolateral reech, along edge. Galeala 15
wm long, tps pointing latcrally, Posterior
hypostomalae 26 pm long, somewhat bulbous
towards base of shafi, shafl with several Jong, surong
setules. Palpal seral formula 1, 6, 3, 4, Palpal
femorala dorsul, spiniform, § «m long. Palpol
tibialae and tarsalae as figured. Palpal tibial chow
With tines robust, subequal but the dorsal the loner
4§
NEW LARVAL TROMBIDIOID MITE FROM AUSTRALIAN CENTIPEDE
100
}
Fig, 1, Wondeclia centipedae gen. nov., sp. nov. Larva, holotype. Dorsal view, entire, shown partly in transparency
(scutal sensillary seta from another specimen). To scale shown; seta “c” is figured in Fig. 3C.
46 R. V. SOUTHCOTT
ae daw
V7 4
a 200
=,
ae
Fig. 2. Wondeclia centipedae gen nov., sp. nov. Larva, holotype. Ventral view, entire, to scale shown; seta “d”
is shown in Fig. 3D.
NEW LARVAL TROMBIDIOID MITE FROM AUSTRALIAN CENTIPEDE
= 0
wo ff
it
>
a
47
Pasctis E
icc Ga
pr
( PaScFe /,.
&
Fig. 3. Wondeclia centipedae gen. nov., sp. nov, Larva, holotype. A Anterior dorsal scutum (sensillary seta is from
another specimen). B Posterior dorsal scutum. C Dorsal idiosomal seta (“c* in Fig. 1). D Ventral idiosomal seta
("d" in Fig. 2), & Gnathosoma, dorsal aspect. F Gnathasoma, ventral aspect. (All to seale shown.)
blunt-pointed; with a slight split.
supracoxalae absent.
Metric data are provided in Table J.
Palpal
Maretial examined: Queensland: Wondecla, in eucalypt
forest, 2,i11,1944, R. V. Southeott,; 30 specimens, (South
Australian Museum): SAM NI9879 (Holotype, also with
label ACB6I3A (RVS)); NI98710-198738 (paratypes):
ectoparasitic upon centipede SAM A391, RAysida nuda
(Newport) (kindly identified by Dr L. E. Koch, Western
Australian Muscum).
Correlations hetween shield and leg variates
In continuation of previous studies (Southcott 1966,
1986a, b, c) into the degree of correlation between
shicld and leg variates in prostigmatic mites, the
data of 15 specimens of I centipedae were
examined by correlalion analysis. The results are
shown in Fig. 7.
Among a tabulation of 49 x 49 variates we may
expect to find a nmumber of significant correlations
by chatice alone; half of these should be positive
and half negative. In Table 2 are shown the findings
for the comparisons, at three levels of probability.
Fig. 7 and Table 2 show that there is a significant
excess of positive correlations, and a deficit of
negative correlations, from random expectations.
There are no negative correlations at the 0,001 level
of probability, and only two at the 0.01 level of
48 R, V, SOUTHCOTT
Fiz. 4 Wondeclia centipedae gen. nvv., sp. nov.
Interpretation and coding of structures and dimensions
af anterior and posterior dorsal scula,
probability, neither of which relates to the same
variate. The negative correlations observed can
therefore be ignored,
The positive correlations appear to be scattered
niore or less evenly through Fig. 7, In Table 3 these
significant positive correlations, over 6 groups of
variates, are examined by a classification into
comparisons between variate groups.
The 1176 correlation coefficients under study are
not independent of each other, but approach
independence ag the sample size increases (see, &g.,
Cameron & Eagleson 1985; also Southcott 1986b).
Table 3.shows that at the 0.05 level of probability
(here is an excess of positive correlations in cach
of the six groups examined — above Lwice int every
one except that of group 4, of shield » leg variates.
Table 3 shows also excesses in the numbers of
povitive correlations at tevels of probability
indicating preater significance, The overall degree
of correlation appears comparable with that
observed in some other trombidivid mites that have
been studied for correlations by similar techniques
(Southcott 1986b), lying between the levels observed
for Trombella cucumifera Southcou and 7. rugosa
Southcott. However, as with the findings in
Trombella, the degree of correladon found is not
sufficiently great to inspire confidence in the use
of a restricted set of variates for species identi-
fications.
Vase 1. Metric data for larvae of Wondeclia centipe-
dace, in wm (except proportions),
Variate Holotype n range mean = sd.
LN 30 15 24-31 27.53 1.9223
MA 27 15 22-27 924,67 1.3452
AW 67 15 57-67 62.40 2.5857
PW 73 15 69-76 72.07 2.1536
SB 55 15 49-55 52.67 1.7593
ASB 49 (5 40-49 = 46.80 2.3664
PSH 19 15 16-20 18,00 1,04K)
L 68 1S 58-68 64.87 2.6690
W 83 15 78-86 =—-B2.80 2.4260
AP 32 15 28-35 = 31.73 1.7914
AM 27 15 22-29 26.33 1.7182
AL 40 [4 32-41 36.29 2.7296
PI. 42 1§ 40-$0 44,20 2.4260
AMB 54 159 47-55 50.13 2.1668
SE — 3. S1-5B) = 54,67 35,5119
LA 3 15 3-5 3.80 0.4761
LP 32 15 8-37) 33.27 7.1461
SA 46 15 38-46 «= 43,53) -2,2949
SP 18 1S 33-18 =15.33 1.3452
PLN 7 is 5-8 6.93 0.9837
PSI. 34 15) (31-35 33,93 1,2799
PSW 7 15 69-81 = 75.33 3.2878
Ow 7 1S 16-20 18.13 1.4573
Ql. 53 15 48-59 54.00 2,976)
RLN 18 16 LS-19) 17,07 1.2228
RW 46 1§ 45-56 49.33 3.4157
RL 35 1§ 49-58 35.13. 2.2949
DS (shortest) 39 15 31-44 39.13 53,2704
DS (longest) 35 15 49-56 52.40 2.0976
MDS 55 15 49-56 52.40 2.0976
PDS 51 1§. 43-5) 47.13 2.4456
Fel 40 1S 34-40) 38.60 1.6818
Gel 29 15 27-29 28.00 0.8452
Til 34 1S 30-36 32.13 1.6847
Tal(L) 62 15 55-63 60.47 2,1336
Tal(El) 7 1$ §5-20 = 16.40) 1.1212
Fell 35 1S 28-36 32.87 2.3563
Gell 25 1S 23-26 24.47 0.8338
Tilt 4] 1S 28-3) 29.13 1.1255
Tall(L) 4 15 50-55 = 52.13 1.7265
Tall(H) 15 1S 18-16 =-15.20. 0.4140
Fell) 40 1S 36-41 37,73 1.5796
Gelll 26 15 22-26 =. 23.60) 1.0556
Till 35 15 29-35 = 31.27) 1.6242
‘TaTI(h.) 59 13) 53-42 S413) 2.2949
‘Tal li{H) 15 15-13-15 14,27 0, 7988
Til/Gel L.U7) US £.03-1.33) 1,15 0.0774
Tilk/Gell L240 US LOB-L.24 1.19 0.0488
TUN /Gei ll 135 (5 L2)-145 132 6.0701
Tans 2 Correlations in variates for larvae of Wondectia
centipedne sp. noy,
P tye correlations ve correlations
Observed Expected* Observed Expected?
0.001 7 1 a i
0.01 34 6 2 6
0.05 113 2y 10 29
“Rounded to whole numbers,
Remarks on the biology of Wondeclia centipedae
Since the centipede host was killed on capture,
few observations on the biology of its mites could
NEW LARVAL TROMBIDIOLD MITE FROM AUSTRALIAN CENTIPEDE 49
jig
Tiny
MLA y f
ae AN 50
m
ne :
Fig. 5 Wandeclia centipedae gen. noy., sp. nov. Larva, holotype. Legs I, I] and [il, to scale shown, to standard
notation, Dorsal aspect of legs I and II, and anteradorsal aspect of leg IIT-
50
R, V. SOUTHCOTT
y
’
af “
oe u
1,!
ty Ti
fi Hy
‘
vr ”
Lal "
rin HY
rity i
wy "
Wa ay i
tT i
Woy t
pier, ~adetates te
YP x
VW
Fig. 6 Wondeclia centipedae gen. nov., sp. nov. Larva, holotype. Legs I, IL and III, to scale shown, to standard
notation. Ventral aspect of legs I and II, and posteroventral aspect of leg IIT.
NEW LARVAL TROMBIDIOID MITE FROM AUSTRALIAN CENTIPEDE
Dontoeciie#0
5]
£snrizedag gen nor., wp. nov.
Ps
2
a
,
5 8B
«ABB
7 PSB
. ot
9 9
w AE Oo } Correlasians
al vy CG
on ee 51 |
i a Oo sextel < other neutat
7 ANB Oo varintes 1964 e +
a 0,04 AY >
ald is Dy OF
G baa is iS) ©
1 ah (ens ) o8 @ 8
190 oaP
ai O O89
u si
2200 (ese Oo
22 OO. Cc
24 QL
25 ORL Oo Oo ia
2h Re
ee . O yo tonomet cotne x othr
zo datubsrtentd O D oo —2__._O % (dionowel getae variates
= tatteegene? (OO 00 0 ra @cutnl = Ldicegaa? a
a a O oe) a0 fe) Oo setas variates fa)
31 Pts. 4} oO a |
so Pet @) ° ie;
Sal Oo a
» TAL
35° Tat(h) qodtel » leg A900 oo
36 tar(e) variates '@; Oo teg 3 other leg
370 Farl CG Tari ates
28 Gell oO © taionomal
39 malt ov Sethe x ing oO
AG TarI{L) oO og varishes
a McIts) o oO
6 oOPaNTT ot a
a3 Gerlt J
as matt 9 o 2)
a> TaITTGL) oO o a ia) ,
360 TaTTI(A)
47 tat /oer © © a] ce} 5
4g TALy/ae1D ie) oO is) oO
49 MITT /GerTt | | o
723 © 5 6 7 B 970 1775293 78-45 1677-78-49 20 21 22 24 20 25 OG 27 GB 29 WO 34 22 33h 35 36 97 38 3G AO AT 8d 45 AH OS AE AP 4B AD
Fig. 7. Correlation matrix of data of Wondeclia centipedae gen. noy., sp. nov., for 49 x 49 variates (see texr).
be made. The dead centipede and its mites were
placed in a tube. Two days later some of the mites
were still alive on the host. No particular sites of
attachment on the centipede were observed.
The mouthparts of the mites are well-developed
for piercing, in fact the cheliccral digits are longer,
more pointed, and with more developed barbs than
in most trombidioid larvae, Presumably the
intersegmental membranes of the centipede would
offer least resistance to puncture,
The absence of eyes in a larval trombidioid is
unusual, and is consistent with a hypothesis that
this larva is well-adapted to a life of parasitism upon
a nocturnal and soil- and litter-inhabiting host. The
flattened idiasoma of the mites may be compared,
for example, with that of the trombiculid mite genus
Babiangia, an ectoparasite upon smooth-skinned
skinks (Southcott 1954). Possibly such flattening
is an adaptation to a mode of life in a restricted
space on a flexible, soil-inhabiting host, and might
reduce the risk of being wiped off as the centipede
moves through soil and litter.
Acknowledgments
] thank Mr L. G. Veitch, Principal Research
Scientist, CSIRO Division of Mathematics and
Statistics, for aid and advice in statistical analysis
$2 R. ¥, SOUTHCOTT
Tan t 3. Classification of positive correlations. between variates in (drvde af Wondeclia centipedae, for observed
and expected values,
Variale groups Numher of
compared comparisons
available
i. Shield » other 351
shield variates
2, Shield xidiosomal 108
setae variates
3, Idiosoma! setac * 6
other idiosomal
setae variates
4, Shickd x leg 486
variates _
‘, Idiosomal setae * 72
leg variates
6. Leg « other leg 153
variates
Totals 1176
Observed numbers Expected
TRUrratyer §
P
0.001 0.01 01.05 0.05
& ig 48 8.78
0] 4 2! 4,7
i i l 0.15
0 " M4 12.48
0 t 7 1k
| 2 il 4.83
8 38 114 29.4
and the Australian Biological Resources. Study for
support.
References
ANDRE, M: (1943) Acariens rencontrés sur des
mynapodes. Bull. Mus. Nat. Hise. Nat, Paris (2)15(4),
181-185.
BerLese, A. (i882) Acari, Myriapoda — et
Psendoscorpiones hucusque in Italia réperta. Fase. |,
No. 4 “Uphis mirabilis Berl. n. sp.", (A. Berlese, Patavii),
(1910a) Lista di nuove specie e nuovi generi di
Acari. Redia 6(2), 242-271.
(1910h) Brevi diagaosi di geneci e specie nuavi di
Acari, Ibid, 346-388.
Cameron, M. A. & EaAqceson, G. kK. (1985) A new
procedure for assessing large correlation matrices. Ausr,
J Slatist. 27(1), 84-95, j
CLOUDSLEY-THOMPSON, J, L. (1968) Spiders, scorpions,
centipedes and mites. (Pergamon Press, London),
(Reprint af 1958 edition).
Domrow, R. {1956} The family Distozerconidar
(Acarina, Mcsostipmata) in Australia. Proc. Lina. Soc.
N.S. 81(2), 193-196.
Evans, G. O: (1955) A review of the laelaptid paraphages
of the Myriapoda with descriptions of three new species
(Acarina: Laelaplidae). Purasitology 45(3 & 4), 352-368.
FAnricius, 5. C. (£775) Systema Entomologiae, sistens
insectorum. classes, ardines, genera, species, adiectis
synonymis,, locis, descriptionibus, observariqnibus.
(Flensburet et Lipsiae).
Leaca, W. E. (1815) A tabular view of the external
characters of 4 classes of animals. Trans. Linn, Sac.
Lond, 11{2), 306-400.
OuDEMANS, A. C. (1912) Die bis jetzt bekannten Larven
van Thrombidiidac und Erythraeidae mii besonderer
Berucksichtigung der fir den Menschen schadlichen
Arten. Zool. Jahrb, Abt. 1, Suppl. XLV, No. 1, 1-230.
Rack, G, (1979) Brennandania scolopendrae sp. 6, von
Scolopendra morsitans (. (Acarina, Tarsonemida,
Fy ppehraraisea, Microdispidae), Kev, Zool, afr. 93(2),
376-381,
Suiba, M, (1976) Taxonomic investigation on free-living
Prostigmata from the Malay Peninsula. Neture and Life
in Saurheast Asia 7, 83-229.
SoutHeort, R. V- (1954) Description of a new genus and
specics of larval trombiculid mite fram New Guinea,
Trans. R. Soc. 8. Aust. 17, 98-202.
(19G1a) Studies on the systematics and biology of
the Erythracoidea (Acarina), with a critical revision of
the genera and subfamilies. Aust. 4. Zool. 9(3), 367-610.
(1961b) Description of two new Australian
Smarididae (Acarina), with remarks on chaetotaxy and
moetaphicat distribution. Trans. R. Soc. 8. Aust. BS,
135-153.
(1963) The Smarididac (Acarina) of North and
Central America and some other countries, Jbid,, 46,
159-245,
(1968) Revision of the genus Charletontia
Hart an (Acarina; Erythraeidae), Aust J Zoal. 14(4),
-819,
(1986a) The genus Odentucurus (Acarina:
‘Trombiculidae). 11. Observations on the life history and
morphology of Odontacarus swan n. sp., and related
forms. Ree S. Anst. Mus. 1912), 169-200,
(19866) Australian larvac of the genus Trombeller
(Acarina: Trombidioidea). Ausé. J. Zool, 34{4), 611-646.
()986c) Studies on the taxonomy and biology of
the sublamily Trombidiinac (Acarina: Trombidiidne),
with @ Critical. revision of the genera. Anat. J Zool.
Suppl. Ser, Na, 123, L6.
(1987) The classification of dhe mite fantilies
Trombellidae and Johisranianidge and related groups,
with the description of a new Jarva (Acurina:
Yrombellidae; Nothro/rombidium) from North
Aincrica.. Trans, R. Soc. §. Aust, 111(1), 25-42.
Tutor, S, (1935) Ubersicht und Einteiling der Familie
‘Trombidildae W, FE. Leach 1814 in Unterfamilien. Zool.
Anz, 95-6), 107-112.
,& Witratann, C. (1947) Trambidiidae. Vierreich
Thb, XxIX-KXXV, I87-S41.
Virzirniim, H, ($940-1943) Acarina. Jn 1H, G. Bron (Ed,)
“Klassen und Ordnungen des Tierreichs”. Band 5,,
Abteilung 4, Buch 5. (Licterung 1-3, pp. 1-480, 1940;
4, pp. 481-640, 1941; §-4, pp. 641-912; 7, pp. 911-10),
1943),
NEW FRESHWATER DIATOM TAXA FROM TROPICAL NORTHERN AUSTRALIA
by DAVID P. THOMAS*
Summary
THomas, D. P. (1987) New freshwater diatom taxa from tropical northern Australia. Trans. R. Soc. S. Aust.
111(1), 53-58, 29 May, 1987.
Two new taxa were discovered during a study of freshwater diatoms from the Alligator Rivers region
of the Northern Territory. Achnanthes pseudohungarica sp. nov. is differentiated from A. hungarica by
the presence of the horseshoe-shaped area on the raphe valve instead of the rapheless valve. Eunotia didyma
var. maxima f. tumida f. nov. varies from the nominate variety by having a consistently smaller length
to breadth ratio.
Both taxa appear to have a limited distribution within the Alligator Rivers region with the Achnanthes
having been found also in the East Finnis River further to the west.
Key Worpbs: Achnanthes; A. pseudohungarica sp. nov.; Eunotia; E. didyma var. maxima f. tumida
f. nov.; tropical Australia; freshwater; Bacillariophyta.
Introduction
The two taxa described herein were found during
a general survey of the freshwater diatoms of
tropical Australia between 1979 and 1981. More
than 160 taxa from 32 genera of diatoms were
identified from the survey (Thomas 1983).
The climate of this region exhibits a typical
tropical alternation between periods of extreme
drought and periods of monsoonal rain. These lead
to rivers varying from a series of disconnected pools
during the “dry” to components of sheets of water,
often several metres deep, which cover thousands
of square kilometres of Australia’s north coast
during the “wet”. Such extreme environmental
variation provides a wide range of growth environ-
ments for algae and a greater regional algal diversity
compared to the more temperate areas of Australia.
Methods
Field samples were preserved as soon as possible
in 10% formalin (usually within an hour). Sub-
samples were then cleared using 50% nitric acid at
60°C for 12 hours (Crawford 1971). The material
was then rinsed of acid by repeated dilutions in
distilled water before being transferred to absolute
ethanol for storage.
Samples were prepared for light microscopy by
evaporating off the ethanol from a suspension of
cells placed on a coverglass at 90°C on a hotplate.
The coverglass was then mounted on a slide using
CAEDEX (MERCK), a synthetic Canada Balsam.
The slides were examined and specimens photo-
graphed with the aid of a Zeiss PM2 photomicro-
scope.
* Botany Department, University of Tasmania, GPO Box
252C, Hobart, Tasmania 7001.
For scanning electron microscopy, coverglasses
with the dried suspension were mounted onto SEM
stubs and sputter coated with gold before being
viewed with the aid of a Philips 505 SEM.
Terminology follows that of Anonymous (1975),
von Stosch (1975) and Ross et al. (1979).
Systematics
Achnanthes pseudohungarica sp. nov.
FIGS 1-5
Valvae lanceolatae, 14-45 ym longae, 5-13 pm
latae. Striae 21-25 in 10 um, ad centrum parallelae,
versus polos leviter radiantes. Areovalva: area
axialis circa 1 pm lata, recta, inter polos secus lineam
mediam formata, saepe crista angusta longitudinali
secus lineam mediam. Raphovalva: area axialis
recta, ad polos circa I wm latos, versus centrum
inflata area 4 wm lata et 7 um longa formans, Area
centralis rhombea, sed area hippocrepica in late
uno. Raphe recta, in lineam mediam areae axialis
formata, spatio 2 um longo ad nodulum centralem.
Description: Valves lanceolate, length 14-45 wm and
width 5-13 pm. Striae 21-25 in 10 um, varying from
parallel at centre to slightly radiate towards the
poles.
Rapheless valve: Axial’ area approximately 1 pm
wide, straight, formed between poles along the
centre line, often with a narrow longitudinal ridge
formed externally along the centre line of the area.
Raphe valve: Axial area straight, varying from
approximately 1 um wide at the poles and inflating
into a central area 4 pm wide and 7 pm long. The
central area is rhomboid but with a horseshoe-
shaped area on one side. The raphe is straight,
DAVID P. THOMAS
-
TROPICAL FRESHWATER DIATOMS 55
formed in the central line of the axial area and with
a 2 wm gap at the central nodule.
Holotype: D930 (D. P. Thomas’ collection,
Tasmanian Herbarium—HO 101052), collected by
K. Bishop on 28.ix.1979 at Magela Falls, N.T.
(12°47'12"S, 133°06'07"E) in slightly acid, freshwater
splashpool at the base of Falls.
Distribution: Also found in Bowerbird, Gulungul
and Nankeen water holes from the Magela Creek
and from Rum Jungle on the East Finnis River,
Northern Territory (see Thomas 1983).
With the aid of the scanning electron microscope
(Fig. 2) it is evident that the cingulum is composed
of the valvocopulae alone and that these are open
and apparently unstructured.
Eunotia didyma var. maxima f. tumida f. nov.
FIGS 6-8
Valvae formam maximam_ similes, sed
abbreviatae et plus inflatae, polis obtusioribus (50°
vs 42°) et ratione longitudinis versus latitudinem
reducta (2.4 vs 3.7). Valvae 76-110 um longae et
38-44 wm latae.
Description: Valves similar to E. didyma_ var.
maxima Hustedt 1913 (Fig. 6) but shorter and more
inflated with less acute poles (50° vs 42°) and a
reduced length to breadth ratio (ca. 2.4 vs 3.7).
Valves range from 76-110 wm long and 38-44 pm
wide.
Holotype: D868 (D. P. Thomas’ collection,
Tasmanian Herbarium—HO 101051) collected by D.
Thomas on 19wiii.1979 at Jim Jim Falls, N.T.
(13°16'34’S, 132°50'12"E) in slightly acid, freshwater
splashpool at the base of Falls.
Distribution: Has not been observed outside the
type locality.
Scanning electron microscopy (Fig. 8) shows that
the valve is ornamented with irregularly arranged,
brief, broad based spines. The cingulum is
composed of a valvocopula and up to three pleurae
with all elements open. The valvocopula has fine
vertical striae (ca. 30 in 10 pm) while the pleurae
appear to be unstructured.
Discussion
Achnanthes pseudohungarica is similar to A,
hungarica (Grunow 1863) Grunow in Cleve &
Grunow (1880) except that the horseshoe-shaped
area is formed on the raphe valve of A, pseudo-
hungarica and on the rapheless valve of A.
hungarica. This is the same feature which was used
to differentiate A. pseudolanceolata Manguin (1962)
(non Hustedt 1952) and A. lanceolata (Brébisson
in Kiitzing 1849) Grunow in Cleve & Grunow (1880).
In addition, A. hungarica has a lower strial density
(16-22 in 10 wm vs 22-25 in 10 wm) and A.
pseudohungarica does not have a narrow stauros
on either valve.
The absence of A. Aungarica from this region
combined with the wide distribution of A.
pseudohungarica make in unlikely that this is just
a mutant variant of the latter and can be considered
a stable species. On the other hand, the very narrow
distribution of Eunotia didyma var. maxima f.
tumida and the presence of its nominate variety in
the same and adjacent water holes makes me very
reluctant to raise it to a higher status than that of
forma.
Acknowledgments
Grateful thanks are extended to Dr Peter Tyler
and the members of the Botany Department,
University of Tasmania who participated in the field
studies upon which this paper is based and made
odd collections in out of the way places. The same
can be said of Mr Keith Bishop from the N.SW.
State Fisheries who collected the type sample of A.
pseudohungarica. The Latin diagnoses were
provided by Dr Tony Orchard, Director of the
Tasmanian Herbarium and the manuscript was
critically read by Dr Tyler. This work was supported
by a grant from the Office of the Supervising
Scientist East Alligator Rivers Region and the use
of their facilities at Jabiru.
References
ANONYMOUS (1975) Proposals for a standardization of
diatom terminology and diagnoses. Nova Hedwigia,
Beih, 53, 523-554.
CLeve, P. T. & GRuNow, A. (1880) Beitraége zur
Kenntniss der arctischen Diatomeen. Kongl. Svenska
Vetens.-Akad. Handl. 17(2), 1-121.
CRAWFORD, R. M. (1971) The fine structure of the
frustule of Melosira varians C, A. Agardh, Br. phycol.
J. 6, 175-186,
Grunow, A. (1863) Uber einige neue und ungeniigend
bekannte Arten und Gattungen von Diatomaceen.
Figs 1-5. Achnanthes pseudohungarica sp. nov. 1. SEM of external surface of the rapheless valve, 2, SEM of external
surface of the raphe valve showing lack of external opening of the horseshoe shaped area and the valvocopula.
3. SEM of internal surface of the raphe valve showing the internal opening of the horseshoe shaped area. 4. LM
of the rapheless valve (Holotype slide HO 101052). 5. LM of raphe valve (Holotype slide HO 101052), All scales 10 um.
DAVID P. THOMAS
TROPICAL FRESHWATER DIATOMS 57
Zweite Folge. Verhandlungen der kais.-k6nigl, zool-bot.
Gesell, Wien 13, 137-162, Plates 13-14.
Hustept, F, (1913) Jn A, Schmidt, et af (1874-1959)
“Atlas der Diatomaceen-kunde”. Plates 285-288. (R.
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(1952) Neue und wenig bekannte Diatomeen. IV.
Bot. Not. 4, 366-410.
KuTzING, F, T. (1849) “Species Algarum”, 922 pp. (F. A.
Brockhaus, Lipsiae).
Manautn, BE, (1962) Contribution a la Conaissance de la
Flore Diatomique de la Nouvelle-Calédonie. Mém. Mus.
Nat. Hist. Nat. nouy, sér, sér. B, Bot. 12(1), 1-40.
Ross, R., Cox, E, J, KARAYEVA, N. I, MANN, D, G.,
Papoock, T. B. B., SIMONSEN, R. & Sims, P, A. (1979)
An amended terminology for the siliceous components
of the diatom cell, Nova Hedwigia, Beih. 64, 513-533.
StoscH, H. A. von (1975) An amended terminology of
the diatom girdle. Nova Hedwigia, Beih, 53, 1-28.
Tuomas, D. P. (1983) A limnological survey of the
Alligator Rivers Region, Northern Territory. 1, Diatoms
(Bacillariophyceae) of the region. Supervising Scientist
for the Alligator Rivers Region, Research Report 3,
1-139,
Figs 6-8, 6. LM of Eunotia didyma var. maxima from sample no. 868. Eunotia didyma var. maxima f. tumida f.
nov. 7. LM of valve (Holotype slide HO 101051), 8. Oblique SEM of dividing frustule showing raphe, girdle structure
and marginal spines. Scale 100 pm.
FEEDING AND GROWTH OF GOLDEN PERCH LARVAE AND FRY
(MACQUARIA AMBZGUA RICHARDSON)
BY P. T. ARUMUGAM & M. C. GEDDES*
Summary
Golden perch larvae were stocked into an earthen pond and grew exponentially from a mean
standard length 4.5 mm (dry weight 0.16 mg) to a mean of 31 mm (165 mg) in 46 days from
12 November until 28 December, 1984. The growth coefficients were 0.04 mm mm’! day for
length and 0.15 mg mg” day” for weight. The mouth gape was related to length. At first feed the
type and size of prey was restricted by poor swimming and pursuit abilities and small mouth gape.
Larvae and fry greater than 10 mm standard length were able to pursue a wide range of
zooplankters but feeding was limited by mouth gape. The daily food consumption of larvae and fry
increased from 33 to 5600 ug dry weight per day. Because of the relatively small size of golden
perch larvae at first feed, survival is dependent upon a high density of appropriate sized
zooplankters.
KEY WORDS: Fish, larvae, fry, Macquaria arnbigua, feeding, growth, mouth gape.
FEEDING AND GROWTH OF GOLDEN PERCH LARVAE AND FRY
(MACQUARIA AMBIGUA RICHARDSON)
by PLT. ARUMUGAM & M. C. GEDDES*
Summary
AKUMUGAM, BT & Gepoes, MC. (1987) Feeding und growth of golden perch larvae and fry (Morquarsa
ambigua Richardson), Yrs. R. Soc. § Aust. V11(1), 59-65, 29 May, 1987.
Golden perch Jarvae were stocked into an earthen pond ahd grew exponentially from a mean standard
length 4.5 mm (dry weighe 0.160ng) to a mean of 31 mm (165 ow) in 46 days from 12 November until
28 December, 9X4, The growth coefficients were 0.04 mm mim-' day! for lengil and 0.15 me mg-! day!
for weight, The manth gape was related to Iength. At Ulyst teed the type and size of prey was restricted
by poor swimming and pursuit abilities and smal! mouth gape, Larvae and try geearer than 1) mor standard
length were able to pursue a wide range of zooplanklers but feeding was limiled by mouth gape, The daily
food consumprion of larvae and fry increased from 33 to 5600 wg dry weight per day. Because of the relatively
4mall size of golden perch larvae at first feed, survival is dependent upon a high density of appropriate
sized zooplankters.
Key Worots; Fish, larvac, Ity, Macguaria emibigua, feeding, growth, mouth gape.
Tniroduction
The golden perch (Macquarta ambigua
Richardson) belongs to the Percichthyidae, a group
ot fishes which includes both freshwater and marine
representatives, Mf, ambigua occurs throughout the
Murray-Darling system excepr.at higher altitudes,
in the Lake Eyre and Bulloo-Bancannia drainage
systems and the Dawson-Fitzroy River system in
eastern Quecnsland (Lake 1971; MacDonald 1978;
Llewellyn & MacDonald 1980; Merrick & Schmida
1984). These references cite the occurrence of M.
embigue in coastal streams in northern New South
Wiles, but there are no self-maintaining populations
in these drainages (Rowland pers, comm.).
Little is known about the biology and ecology
of golden perch larvae and fry in the natural
environment. An upstream spawning migration 1s
tnitiated. by water level rises at the onset of major
floods (Reynolds 1983), The semi-buoyant cegs are
probably carried downstream by the flood waters
with the larvae hatching and drifting in the water,
before entering shallow floadplain areas and
“billabongs", which probably act as nursery grounds
(cl. Lake 1967a; Rowland 1983; Rowland pery.
coinm.). These Floodplain areas receive nutrients
and allochthonous materials that promote increases
in plankton and other organisms essential for
survival of (he larvae and fry (cf. Shiel 1980; Maher
1984; Briges ef a/, 1985), In many areas dams and
weirs have blocked spawning migrations, lowered
water temperatures by discharging colder water
from the deeper water layers of dams and reduced
* Department of Zoology, University uf Adelaide, G.B.O,
Box 498, Adelaide, S. Aust, SOOL,
flooding which is necessary to induce spawning and
provide suitable conditions for the pelagic eges and
space and planktonic toad for the young tish (Lake
197); Reynolds 1976; Cadwallader 1978; Pollard e¢
al. 1980; A.R.LE.R. 1983).
Much of the available information on golden
perch larvae and fry comes from research into
spawning and larval rearing carried out at the In-
land Fishcries Research Station (LF.R.S,}, Narran-
dera (Lake 1967a, b; Rowland 1983, L9&6a, br
Rowland ef ei, 1983), Golden perch are induced to
Spawn using humal chorionic gonadotrophin and
the larvae are reared in earthen ponds trom first
feed until they are aboul 25 to 30 mmm standard
length, usually a period of 25 to 35 days, Lake
(1967b) observed that the stomach contents of lar-
vae at first feed consisted of cladocerans, copepods
and phytoplankton.
The present study was conducted at the I,F.R,S.
The main objectives of chis study were to describe
the growth characteristics of the golden perch lar-
vac jn rearing ponds, to investigate the behaviour
and ecology of the larvae and fry in relation to feed-
ing, and to measure daily food consumption of the
different size larvae and fry, The description of
growth involved the determination of length-weight
relationships and the estimation of the growth rate
of golden perch in a rearing pond, Particular em-
phasis was placed on the relationship berween
length and mouth gane, as mouth gape determines
the size of prey that wan be taken (Shirota 1970),
This information, In conjunction with that on feed-
ing behaviour and daily food consumptian will pra-
vide useful guidelines to the conditions required far
survival and growth i golden perch larvae.
a P, T. ARUMLIGAM 4 M. ©. GEDDES
Methods und Materials
Growth Characteristics
Larvae and try were caught froin a rearing pond
ar LER.S. over a period of 46 days trom 42
Novernber-28 December, 1984, They were killed
with an overdase of soda water, standard and rotal
lengths were measured, wet Weights were determined
and then they were dried in an oven fer 24 hours
at 40°C, and dry weights measured. To establish the
power relationships between weight and length,
Jinear fegressions were performed on the log
transformed data of both lengths and weights
(Hagenal & ‘Tesch 1978). For the total length-
standard leneth and dry weighi-wel weight
relationships, regressions were performed an
untranstormed data. The gruwih rates of the larvae
and fry in the pond were analysed hy establishing
standard length-time and dry weight-time
relationships, An exponential model gave The best
fit und so both standard length and dry weigh!
méusuremenis were log transformed. Mouth gape
and standard length were measured for about 450)
specimens collected during 1983 and 1984 and
preserved with forinalin. ‘The mouth gape was
measured as the external distance bel ween the two
corners of te mouth when shut. The regression of
mouth gape and standard length was perlormed
witherut transformation.
Behaviour
Observatiuns ow feeding and cercitoriality were
tmade on fry from first feed to abour 30 mm
sandard lenotl) either alone in 250 ml capacity
beakers or in groups of 10-15 in 70 | aquaria, The
larvae were ted with brine shrimp (Artemia), using
(he nauplil for ybe smaller larvae and adults for the
larcer larvae and fry. Zooplankters and chironomid
larvae were occasionally fed to the larvae and fry.
Cover was provided im the form of a 30-40 mm
length of opaque, plastic tubing with 2.5 mm
infernal diameter for the smaller larvae and a fl 70
mm length of tube with imernal diameter of 3,0 mm
lor the larger fry (25-30 mm). One cover was
provided for the fry held singly and che number of
covers was ahout two-thitds the number of fry for
Lie group surdy, The study was carried our at
1) 24°C under daylight condijions,
Daily foud consumption
The daily food consumption of u fish, estimated
as the number of food items supplied over 24 hours
minus the nurnber ol ems remaining (cf Gophen
1980), Was estimated for size classes of 4.5 (= first
feed), 10, 20 add 30 mm standard length, Treshly
hatched brine shrimp nauptii (length: 400-600 ym)
were used as food except for che 30 mim fry where
Daphnia carinata (enyth: 100-2320 pm) were used
because of {he enormous number of 6rine shrimp
nauplii that would have been required. The leeding
regime involved sinall numbers of food items being
fed to the larvae and (ry al a time, rhe next feed
being given when only a few food items remained.
Three to six feeds were required over the 24-hour
period. Por the feeding trials, five different
individuals of . particular size were placed
simultaneously in separate beakers which received
circulating filtered pond water at 20-22°C. Phe fry
were no} slurved prior to commencement of feeding
trials and a tuhe was provided as 4 réluge for the
fry. ‘Uhe daily food consumplion was expressed as
nuunber of ems and as pe dry weight consumed
per day, Dry weights of brine shrimp nauplil and
Daphnia were obtained from weighing a known
number of oven-dried specimens al 60°C and
obtaining the weight per individual brine shrimp
nauplius (2.55 yg) or Daphnia (20 we).
Results
Larvae al first feed were 4.4-4.6 mm standard
length (mean 4,5 mm) (total length - 4.5-4.7 min:
mean 4,6 mm) with a dry weight of 0.14-0,18 me
(mean 0,16 mo} (wet weight = 0.47-0,53 mg; mean
0.52 meg). Their mouth gape at first Teed was @.5
mm, After 46 days in the rearing pond, they reached
30-31.6 mm (mean 30,6 mm) standard length with
a dry weight of 154-195 ma (mean 165 meg) (wel
weight — 667-743 mg; mean 69L meg), The
regression equations deserihing Jength-weieh|
relationships, che relationship between length and
mouth gape and the growth rate of Try and their
correlation coefficients are given in Table t. The
correlation coefficients ol all pairs of relationships
are highly significant. In the length weight
talationships, regression coefficients (- slopes)
ranged from 3,29-3,72 and for this range growth
is cousidered allometric (cf, Bagenal & Tesch 1978).
The standard length of the fry was directly
Proportional to the total length and dey weight was
linearly related to wet weight, The exponential
growth in beth srandard Tength and dry weight are
shown in Fig. f. The growth ovefficients for
standard Length and dry weight were 0.04 mm/mm
day and 0.55 rag/my day tespectively (Table 1), The
mouth gape of the try was directly proportional to
their standard length with a slope of 0.13 (Fie, 2),
The larvae at First Teed swam [reely in the water
column. They exhibited an innate feeding behaviour
of dacting forward and gulping even when no food
particles. were present, Aller two to Ultree days of
fecding,, larvae could follow and capture brine
shrimp nauplii. efficiency in capturing Mota was
lower, When the golden perch were 10 min aud
FEEDING AND GROWTH IN YOUNG GOLDEN PERCH él
TABLE |. Regression equations, intercepts (a), slopes (b).and correlation coefficients (r) for weight-length, standard
length-iolal length, wet weight-dry weight, mouth gape-standard length, dry weight-time and length-time relationships
in golden perch. (NW = number of data pairs; CL = 95% Contidence Interval; P>0.01; DW = dry weight (mg);
WW = wet weight (mg); SL = standard length (mm); TL. = total length (mm); MG - mouth gape (mm); T =
time (days); € <= exponential),
Y x N Equation a b t
(+/- 95% CL)
DW SL 29 Y =a 7.5 x 10-4 3,61 O.997*#+
(0.10)
DW TL 29 Y =a Xb Lt x» 10 4 3,29 0.998%"
(0,08)
Www SL 29 ¥Y =a Xb 2,9 x 10-3 3.72 U,9ugrr"
(0.18)
Www TL 29 ¥=ake 3.8 x 10-3 3.39 0.995*"*
(0.13)
WwW DW 29 Y=a+dxX 7.98 4.09 0.996***
(0.14)
SL TL 29 Y-a+bxX 0,78 0.79 0.999% **
(0,01)
MG SL 415 Y=ai+bxX 0.23 0.13 0.984*+*
(0.10)
DW T 9 Y = a cbs 0.22 O15 0.99) ***
(0.10)
SL T 9 Y=ae 4,81 0.04 0,992+**
(0.0046)
not have cover stayed in the water colurnn or at the
304 1 bottom of the container away from “defined”
lerritories. When food supply was low, the fry in
covers were healthier and had more convex bellies
van than fry without cover, and with continued low food
= Pa availability mortalities occurred first in fry without
= 2 e cover. Larvae and [ry of 10 mm and greater could
5 53 pursue cladocerans and copepods with ease but the
2 = size of prey engulfed was limited by their mouth
8 f = gape. They altacked very laree Daphnia but released
a them as they were too big ta swallow. (ndividuals
inf } , could slowly engulf a chironomid larva width-wise
? until the whole larva was swallowed.
The larvae at first feed, ate 7-21 brine shrimp
ia 4 5 ell -s nauplii per day (mean 13), 10 mm larvae ate 240-251
Fy 25
-Number of Days Number of Days
Fig. |. The growth in length and weight of golden perch
larvae and fry from release into the pond on 12
November until harvest on 28 December 1984, The
points represent mean values from 3 to § fish on each
sample date.
greater, they preferred cover when kept alone or in
groups and exhibited territoriality. They stayed
inside their tube, with part of their head up to the
eye level protruding, coming out to feed or lo chase
other larvae or fry away, When disturbed, they
would temporarily retreat inside the tube. When a
larger tube was provided, two to three fry sometimes
shared it. In group experiments those fry that did
nauplii (mean 247), while 20 mm fry ate 790-1631
nauplii per day (mean 1110). The 30 mm fry ate
141-423 Daphnia per day (mean 280). The daily
food consumption of larvae and fry ranged from
33-5600 pe dry weight per day and was directly
proportional to the standard length of the fish
(r 0,899), (p< 0.05%) (Fig. 3). There was a large
individual variation in daily food consumption as
indicated by the spread of the points on the graph,
Discussion
Size at first feed is a critical feature of the biology
of larval fish but there is only limited information
osthe size at first feed of Australian freshwater fish
(Table 2). It is clear that golden perch, along with
two other large Australian freshwater fish that lay
62 P, T. ARUMUGAM & M. C. GEDDES
Mouth Gape (mm)
(0 20 30 ao 50 60
Fish Standard Length (mm)
Tig. 2. The relationship between standard length and
mouth gape for golden perch larvae and fry from first
feed to 50 mim standard length.
small pelagic eggs, silver perch (Bidyanus bidyanus)
and silver barramundi (Lates calcarifer), possess
larvae that aré particularly small at the time of first
feed. By comparison the larvae of fish such as
Murray cod (Maccullochella peeli) and freshwater
callish (Jandanus tandanus) that lay demersal eggs
are up to three times longer than M. ambigua larvae,
while those of buccal incubating saratoga
(Scleropages feichardti) and fork tailed catfish
(Arius graeffei) are up to seven times longer than
golden perch larvae at first. feed. The small size of
the golden perch larvae presents problems for
feeding and survival similar to those faced by the
10000
8000
6000
40m .
200
Daily Consumption (yg) doy!)
30 40
20
Fish Length (mm}
Fig. 3. The daily food consumption (pg dry wt day -!) of
golden perch larvae and fry held in the laboratory and
fed brine shrimp nauplii or Daphnia.
larvae of marine fishes during the “critical period”
of larval mortality (May 1974), Another percicthyid
that breeds in fresh water, the North American
striped bass Merone saxatilis, has larvae that are
only 3.1 mm at hatching and 6-7 mm at the time
of first feed and face survival and [eeding problems
similar to those of golden perch (Doroshav 1970).
The allometric growth form of the fry indicates
that golden perch change their body form as they
grow. This is consistent with the observation that
golden perch fry acquire adult features very early
in their development from when the fry are about
11 mm total length (Llewellyn & MacDonald 1980),
TABLE 2, Total length (TL in nmi), size of mouth gape (MG in mm) at first feed and initial food ttems. af the larvae
of some Australian Jishes.
small copepods/cladaccrans
(<500 microns), algac.
mainly small] Moina/ Daphnia
chironomid larvae,
Food. items References
Lake (1967b),
Rowland (1986a)
This study
Lake (1967h)
Daphnia, copepods
mainly filarnentous algae/
Lake (1967b)
phytoplankton
mainly filamentous algae/rotifvers
plankton
microcrustaceans, insect
Arumugam (unpubl)
Lake (1967b)
Rimmer (1985)
larvae, filamentous algae
Species TI. MG
Macquaria ambigua 5.5 —
4,7-4,9 0,5
Maccullockella peeli 12 —
Bidyanus bidvanus 5.5 _—
4.6-5.4 0.4
Tandanus landanus 13 _
Arius grae/fet 50-59 _
Seleropages leicherdti 35 =
Lates calcartfer 3.5-4.5 _
Cyprinus carpio 6.4 0.55
Perea fluviatilis
_ algae, ciliates
rotifers/algue
smal! cladocerans/copepods
= Lake (1971)
— Moore (1982)
M. Mackinnon
(pers, cornm,)
B. Pierce (pers. commi,)
= Lake (1967b)
Guma'a (1978b)
rotifers, cyclopoid nauplii
FEEDING AND GROWTH IN YOUNG GOLDEN PERCH it
Data from I.E R.S. shows that metamorphosis from
the larva to the fry stage is completed when fish
ure about [5-18 mm, 20-29 days atter spawning
(Rowland pers. comm.).
The growth curves. for both length and weight of
the golden perch in the present study were best
described by an exponential relationship with time.
The exponential curve also fitted both length-fime
and weight-time regressions for the first growth
§tanza ota wild population of Perca fluviatilis fry
(Guma‘a 1978b) but other workers have found that
while growth by weight might he exponential, length
inereases linearly, Thus Swanson & Ward (1985)
found that the best fit for total length-time
Tepressioiy curve for walleye (Stizostedion vitreum
vitrenin) was achieved with untransformed data and
A.R.LE-R. (1983) assumed linear Jength-time rela-
Lonships for common carp (Cyprinus carpio),
goldfish (Carassius aurarus) and redfin. However
in these studies, the sampling intervals were so far
apart, especially in the earlier larval phase, thal any
possible exponential relationship during the early
growth phase would have been obscured. The
exponential rate of growth of golden perch larvae
and fry (0,15 mg mg! day -!) can be compared to
the growth rate of fish fry and juveniles, expressed
as Yo increase in weight per day, reviewed by Brett
(1979). Growih cules varied from less than 1% to
Dophnia
Lhe
Boeckella
th {
" Mona
Simm
fish 1Omm fish
— en
MOUTH GAPE OF FISH
20 mm fish
30mm fish
as high as 23% tor small (5 mm) Cyprinodon
macularis, Most figures for various fish under good
conditions were between 1 and 5%. The value of
13'% for golden perch in the present study represents
relatively a high rate of exponential grawth. This
is not surprising as the golden perch were very small
and were in a pond that was fertilized and managed
to promote maximum survival and growth.
The mouth gape of the larvae at first feed is
important because it determines the size of food
items {hat can be taken during this critical period.
Shirota (1970) showed that mouth size of marine
fish larvac at first feed ranged from.200 to 1000 um
and that the range of prey was related 10 mouth size.
In many fish, the size of prey eaten has been shown
to change with growth and the associated increase
in mouth gape (Shelbourne 1962; Einsele 1965;
Shirota (970; Wong & Ward 1972; Siefert 1972;
Guma'a 1978a; Townsend 1983). Therefore
comparisons between the mouth Bapes of different
length golden perch larvae and fry and the array
of common. zooplankters in the rearing ponds will
indicate the food items that can be taken by
different sized fish (Fig..4), The larvae at first feed
are smaller than large Daphnia curinata and
chironomid larvae. Their mouth gapes are similar
to or larger than the smallest Daphnia, small
Mojna, copepodites, cyclopoids, copepod nauplii
chironomid
cyclopoid
golden perch larva
rotifers "s
Fig. 4. Relative sizes-of the development stages of some common zooplankiers and insect larvae in the fry rearing
pond, golden perch larva at first feed, and the mouth papes ol golden perch larvae and fry of different standard lengths.
ed PT. ARUMUGAM & M. C. GEDDES
and rotifers, making them all potential prey. The
potential prey size jicreases asthe try grow. At 30
mm standard lengh, the mouth gape of golden
perch is similar to of larger than the lareest Duphniae
available in the plankton.
The observations on fecding bebaviour suggest
thal goklen perch farvae at first Feed would have
a telatively poor capture efficiency, For striped bass
at firse feed the estimared strike efficiency on
Ariemia nauplii was only about 2% (Miller 1977
in Setzler er al. 1980), More mobile zooplankters
would be less prone tn being captured. Cladocerans
are more sluggish and miore conspicuous than other
xooplankters such as copepods and would be more
prone to. being eaten by the larvae (Zaret 1980). The
feeding efficiency of the golden perch larvae
improved very rapidly and after 2 ta 3 days they
were able to follow slow-moving brine shrimp
nauplii and capture them with ease. At 10mm, they
were agile and last enough © allack any
cladncerans, copepods, ar chironomid larvae, Being
a faster swimmer with good escape responses (Zarer
1972) would not be an elective stralegy for a prey
at this stage. The specd and agility of che [ry meant
Uial once they Visually sighted their prey, capture
was inevilable, In Fact, zooplankton that are faster
swimmers and cover a preater distance (eg,
copepods) may become more prone tu being
predated because the prebability of encountering
the fish would be greater (Townsend 1983),
Although there was a high correlation between
daily food consumption and standard length of the
ry, variability of datly food consumption for the
larger fry was high, Fry used in feeding trials were
individuals recently caught from the pond and so
different growth histories and different levels of
sallauon may have contributed to high variability
(Fuliow & Persson 1978), Also, rhe larger [ry were
easily disturbed and this may have affected feeding.
Considering thal the food intake increayed as a
linear Function of size and that growth of golden
pereh in the pond was caponential with lime, the
daily food consumption would also inerease
exponentially with time. Thus the intensity of
predation by the fry on the zooplankten
hopulations Would increase slowly at first but at
later stages it would increase rapidly and migtt
produce @ sudden impact on the zooplankton
community.
The information on feeding behaviour, growth
and the mouth gape, fighlight (he difficulties faced
by golden perch larvae and Fry, Survival and grewth
requires an environmeat with a high densiry of
appropriate sized food su that larvae with limited
feeding ability and mouth gape can predate
successfully. Mouth gape increases only slowly
during the first weeks and so the range of
zooplankters available to the fry will be limited for
some lime und high densities of small {<1 mim)
zooplankters will be necessary for high survival and
growth rates, These requirements are met at the
LER.S. wheré ponds aré Flooded just prior to the
release of fish fry and the early stages of
zooplankton succession are dominated by rotifers,
small cladocerans (especially Maina) and copepod
nauplii (Arumugam & Geddes 1986), In the wild,
the same conditions may be provided an recently
inundated floodplain areas where zooplankton
communities are in early successional srages. Lf so,
then the possession of pelagic eggs and the timing
of breeding to coincide with rising water levels may
allow golden perch larvae to be dispersed over
recently inundaled floodplains. where they find
conditions that are suitable fur their development.
The availability of cover for the {ry in these areas
may also affect the grawth and survival,
Acknowledgments
We thank the Department of Agriculture, New
South Wales, for permission fo carry out the study
at the Inland Fisheries Research Station.
Narrandera, Thanks tothe LF.R.S. sratf, especially
Stuart Rowland, Steve Thursian and Iohn Dirou
for their help and advice, We alsa thank Stuart
Rowland for critically readirig the manuscript. The
work Was made possible by the provision of a Post
Graduate Fellowship by IDP to one of us (PTLA.)
and support froin FIRTA. Thanks to David Culver
for assistance in the field, Ruth Evans for artwork
and Sandra Lawson and Heather Kimber for word
processing several drafts of the manuscript,
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perch. Muequaria ambiquu (Richardson)
(Percichthyidae). Ayuacnliure 35, 221-238.
(t986a) The hormone-induced spawning and larval
redring of Australian native Treshwater fish, wilh
particular emphasis on the golden perch, Macquaria
amingua, Proceedings af the First Freshwater
Aquaculture Warkshop, Narrandera, NSM, 21-25
February 1983, (Department af Agriculture, Sydney).
(1986b) Design and operation of an extensive
aquaculture system for breeding warmwater fishes, [bic
, Dirou, J, & SeLosse, P, (1983) Production and
stocking of golden and silver perch in N.SW, Aust. Phish,
42, 24-28,
Skivter, BE. M,, Bayntom, W. Ru Woop, K. V., ZION,
H. Hy, Luppers, L., MOUNTFORD, N. K., Frere. PB,
Tucker, L. & Mtnursky, J. A. (1980) Synopsis of
biological data on striped bass, Marone saxatiffs
(Watbaum). FAO Synopsis No, 121. 69 pp.
Siecnougwp, J. B. (1942) A predator-prey size
relationship tor plaice larvae feeding on Orkopleura
fi Mar. Mial, Assoc. UK. 42, 243-252.
SHiEY, R. J. (1980) Billabongs of the Murray-Darling
system, pp. 376-390, In W. D, Williams (Ed) “An
Feoloaical Basis for Water Resource Management,”
(Australian National University Press, Canberra)
SHinuta, A, (1970) Studies on the mouth size of lish
lorvae, Bwll, Jap, See, Se, Fixh. 36, 353-368,
Sievert, R. B. (1972) First food of larval yellow perch,
white sucker, bluegill, emerald shiner and ralnhow sarelt,
Trans. Amer. Fish Sac, 101, 219-225,
Swanson, G, M. & Wake, F. J. (1985) Growth of juvenile
multeye, Stizostedion vitreunt vitreunt (Mitchell) in twa
man-made ponds m Winnipeg, Canada, Verh. Int. Ver.
Limwol, 22, 2502-2507.
‘Townsenp, BD. W. (1983) The relations between farval
fishes and zooplankton in two inshore areas of the Gulf
of Maine. 4 Plankton Res, 5, 145-173,
Wona, RB. & Warn, F. J. (1972) Size selection of Duphniu
pulicariu by yellow belly (Perca flayescerns) fry in West
Blue Lake. J. Fisk Res. Bd. Canada 29, 176)-1764,
FARET, ‘VM, (1972) Predator-prey interaction in a
tropical lacustrine ceosystem. Evology $3, 248-257.
(1980) “Predation and Freshwaler Communities”
(Yale University Press, New Haven and London), 187 pm
TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL, iil, PART 2
PESTICIDE CONCENTRATIONS IN SOME SOUTH AUSTRALIAN BIRDS
AND OTHER FAUNA
BY P. R. BIRKS* & A. M. OLSEN}
Summary
Data are provided for concentrations of the pesticides DDT plus metabolites, HCB, lindane, aldrin
and dieldrin present in tissues of 20 native and two exotic bird species, one freshwater crustacean,
one amphibian, two reptiles (lizards) and two exotic mammals from Saddleworth-Riverton survey,
1972; 18 native and two exotic bird species from miscellaneous areas, 1968-74; eggs of 23 native
and two exotic bird species from miscellaneous areas 1971; and 26 foxes from miscellaneous areas,
1973.
KEY WORDS: Organochlorine pesticides, Aves, Reptilia Amphibia, Crustacea, exotic Mammalia,
South Australia.
PESTICIDE CONCENTRATIONS IN SOME SOUTH AUSTRALIAN BIRDS
AND OTHER FAUNA
by P. R. Rirks* & A.M, OLSENT
Summary
Hinks, BR, & Orsie, A.M, (1987). Pesticide conventrations in some South Australian birds and other
tauna. Zeans. KR. Sov. S. Aust. 11(2),.67-77, 29 May, (987.
Data aré provided for concentrations of the pesticides DDT plus metabolues, HCG, lindane, aldrin
and dieldrin present in Ussues of 20 native and two exotic bird species, one freshwater crustacean, one
amphibian, (wo reptiles (lizards) and two exorie mammals from Saddleworth-Riverton survey, 1972; 18 mative
and two exotic bird species from miscellaneous areas, 1968-74; eggs of 23 native and nyo exotic bird species
from miseellaneous aveas 1971; und 26 foxes from miscellaneous areas, }973,
Key Wokbs: Organacblorine pesticides, Aves, Reptilia Amphibia, Crustacea, exotic Mammalia, South
Ansatralia-.
Introduction
There is a paucity of published data on pesticide
contamination in Australian fauna. In Victoria, 24
widely Separated areas, each covering about 9300
ha, were surveyed for determination of
concentrations of pesticides in 63 bird species, 10
mammal species, 15 fish species and eggs of 14
different birds (Butcher 1967; Australian Academy
of Science 1972). In the Nucthern Territory Best
(1973) surveyed the organochlorine pesticide
residues in the fatty tissues of 12 mammal, four
bird, ten reptile and six fish species from
undeveloped and develaped areas in arid and
tropical zones.
The Australian Academy of Science (1972)
reported ranges in concentrations of DDT plus
metabolites in (he Namoi cotton growing region for
waters of creeks and rivers and their sediments,
algae, crustaceans, fish, birds and turtles together
with the range in concentrations of 29 fat samples
from ten species of birds of the Bathurst area.
Olsen & Settle (1979) when reporting on the
pesticide contamination in various tissues of water
rars in the Murrumbidgee Jrrigation Area, N.S.W.
1970-72 indicated that “little is known of the
pesticide contamination of Australian fauna”.
The South Australia Pesticides Advisory
Cununittee was concerned about the lack of data.
on the concentrations of pesticide residues in fauna
and the implications of this for research planoing.
AS a consequence, in mid 1971, programmes to
obtain these much needed baseline data were
commenced, The Department of Fisheries and
Fauna Conservation arranged for eggs of a number
of bird species to be collected in the spring of 1971
* Dept of Agriculture, 25 Grenfell St, Adelaide.
§. Aust, 5000.
[1 Orchard Grove. Newrm. & Ause 0/4,
while officers of the Deparcment of Agriculture
undertook the collection of representative faurta in
a selected arcu in January 1972.
Broad surveys 10 determine concentrations ot
pesticides in native and exotic avi-fauna and some
other terrestial fauna from selected areas in South
Australia were undertaken between 197] and 1973.
Determinations of pesticide residues in hirds
suspected of pesticide poisoning were carried out
between 1968 and 1974,
History of Pesticide use —
Saddieworth-Riverton arca
po’ DDT
This area was selected because of the relatively
high DDT usage for control of pea weevil (Bruchus
pisorum) and native badworm (Heliothis punctiger)
in field pea crops, As it is usual to have a 7-8 year
crop rotation and the average annual area sown Lo
peas is about 3%, it follows that abour 20% of the
total area of aver SOM) ha would have received direct
applications of about 1.5 kg/ha of pp'DDT in the
8-year period prior to 1971. The area has an anual
rainfall of about 500 mm.
In October 1971, 150 hectares of field peas were
treated with pp'DDT; 40 ha were sprayed with 0.7
ke/ha pp DDT in early October and a second
spraying, ai the same cohcentration was giver later
in the month whilst the 109 ha paddock of peas was
sprayed once with 1,05 ka/ha pp'DDT. Thus a total
of approx. 170 ke of pp'DDYT was applied over 150
ha three months before the fauna sampling study
was commenced in late January 1972, Lucerne
growiog In the area was not sprayed.
Lindane
The only report of the use of lindane in the
SaddJeworth-Riverton atea was a3 lindane-
OLSEN
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IDES IN SOUTH AUSTRALIAN FAUNA
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superphosphate for the control of redicgged
earthmite (Halatyeeus destructor) and lor pests of
germinating cereals in May to July 1971.
Dieldrin
The use of this organochlorine compound tor
pasture or crop spraying was discontinued In the
Saddleworth-Riverton area in 1967,
HCB
The only wse of FICB was as a seed dressing and
its use and availability ceased in 1972_
Methods
All bird and mammal specimens in the
Suddleworth-Rivetion area were Laken by shooting
with 0410 gauge shotgun or 0.22 calibre rifle
Whereas the lizafds, frogs and yabbies were taken
alive, All Specimens showed normal behaviour al
the time of collection.
All dowd specimens were placed individually in
polythene bays approved by the Chemistry Division,
Deparhmeat of Services and Supply, The specimens
were labelled and packed in ice lor delivery to
Department of Fisheries and Fauna Conservation
for identification checks before being sent for
analysis by gas liquid chromatugranhy in the
Chemistry Division, Limits of detection were HCB
U.007 me/ke, lindane 0.01 mg/kg, dieldrin (01
me/ky, PDE 0.01 mg/kg, DDD 0.05 me/kg,
pp’ DPT 0,07 mu/ke,
Some bird specimens were badly damaged by
shat and for that redson only the available parts
of the bodies were used in the analysis. The wines
and feathers of all birds were removed before
extraclion,
Some data on pesticides residues in a few native
birds trom miscellaneous areas taken incidentally
in the course of other studies as well as results of
analyses of birds forwarded for examination by
interested people suspecting that the birds may have
died from pesticide poisoning were obtained,
A limited sampling of birds’ cgas for pp'DDT
plus metabulites was undertaken in the spring of
1971, Single eggs Irom chitches of two or more eges
were collected from a wide range of habitats
extending from Lyndhurst in the north of the State
to Lucindale in the south-east by AH. 3, Morton of
Jervois. No shell thickness measurements were
ntade.
During 1973 the Animal Health ranch, Depart-
ment of Agriculture, curried out a survey of the
imestinal parasites of foxes, the animals being taken
trom widely scattered districts Uhroughqut the State.
The Pesticides Commitiee requesied that samples
of fac adhering to the Kidney be taken for pesticide
wtralysis.
Results
Saddleworth-Riverian area
Table | shows indivMidiial pesticide concentrations
in the Taunt examined from (he Sadedlewerth-
Riverton area.
The metabolite DDE was present in all birds
while the parent pp'DDT was present at low
concentritions or absent,
The rearrangement of the data in descending
order of DDT plus metabolites (Table 2) indicates
thar pesticide coricentrations were extremely
variable, the highest and lowest were in grain feeding
ypecies. However, the high concentration in the
house sparrow (a grain feeder) and the lower levels
in the laughing Kookaburra and the Australian
kestrel (meat-ealers) demonstrate the need for mare
data before a relationship can be postulated between
residues and diet.
In thus! cases, those spevies which are known
from data of the Australian Bird Banding Scheme
lo show least mayement From banding sites, were
those which had the higher concentrations af
pesticides in their tissues.
Miscellaneous areas
The analyses of 35 birds representing 21 spectes
from miscellaneous areas (Table 3) showed that 14
birds had concentrations above 1 meg/ke DDT plus
metabolites, (wo birds had above 0.5 but less than
10 mg/kg and 19 birds had less than 0.5 mg/ko
in various body rissues.
The two highest concentrations were in birds
forwarded by concerned people who considered that
the birds may have died froin pesticide poisoning,
A grey bulcherbird (28) mg/kg), which came [rom
afarm on Eyre Peninsula, is believed to have died
from ealing dield mice froma grain Mere which had
heen dusted with pp'DDT for rodent control during
the 1970 mouse plague. Even then the use of
pp’ DDT as a rodenticide was nolalticially approved
or recommended. likewise ihe sucred kinglisher
from Rendeclshim (South-East) with a
concentration of 75.6 me/ke DDT plus metabolires,
Was suspected by the finder, of poisoning from
pesticides.
The next highest congenfrations were in an
Australian pelican (2.4) mye kek a little penguin (5.2
mg/kg); a stubble quail (4.2 mg/ks); another two
fitthe penguins (4.0 atid 3.8 mg/ke respectively) ana
a darier (3.78 mg/kg) all of which were collected
opportunistically, Four young little penguins came
Ashore with oi! soaked bodies: they were starving,
Because of the seareity of thelr body fat, oil from
the preening gland was used in the analyses,
itis doubtful whether the presence of pp’ DDT
was tcspousible for the deaths of birds from home
PRSTICIDES IN SOUTH AUSTRALIAN FAUNA 7
TABLE 2. Descending order af concentrations uf DDT plus nietaholites in birds — Saddleworth-Kiverton area 1972
Prime-Food*
Small
Animals
Species
Invert-
ebrates
House sparrow
Magpie-lark 4
Australian magpie + {
Willie wagtail +
White-faced heron + i
Black faced wood-
swallow ’ hocy
Striated pardalote '
Little raven ' {
Noisy miner \
Brown treecreeper \
Hooded robin \
White-plumed
haneyedlec ‘
Red-rumped parrot
Brown-headed
honeyeater '
White-hrowed
swallow '
Laughing kookaburra \ ‘
Galak
Galah
Australian kesirel ' +
feral. piecon
White-winged chough +
Cockatiel
Crested pigeon
Concentration (mg/kg — wet weight)
Seeds Fat
Fruits Max Other Tissuc
* 282.3 body
‘ 148.8 liver
\ 95.8 lives,
25.3 stomach
61.9 16,2 body(2)
46.7 liver
43.8
+? 39.0 15.2, 8.0 body(3)
+ 39.0 stomach
+7? 33.2 16.3, 14.5 hody(3)
{8.8 9.4 body(2)
16.2 bady
13,2 5.5 hody(2)
. 6 3.98, 0.67, body(6)
0.6, (1.46, }
0.44
8.1 bady
8.0 body
7.68 stomach
+ 6.0 liver
4.7 stomach
3.94 liver
+ 2.47 liver
+ 2.1 14 liver(2)
+ 1s t.16, 0.33 body(3)
+ 0.34 liver
(after Lim 1972 see Table [)
* Invertebrates = insects, crustaceans, and some mollus¢s. Small animals = fish, frogs, lizards and young birds.
Seeds and fruits = includes bulbs and such like underground plant storage tissues.
gardens in various areas of the State (Table 3) when
apparently healthy birds of the same species with
much higher concentrations of the pesticide were
surviving without showing any stress symptoms
(Table 2). In post-mortem examinations of the
cormorants and the pelican by veterinarians, death
was ascribed to respiratory infections.
Twenty-six cggs of 23 species (21 native species)
were examined and no evidence of pp'DDT or
metabolites was found in epgs from eight species.
Atiother eight species had a range of concentrations
from 0.01 to 0.10 mg/kg DDT plus metabolites,
whereas seven species had egys with concentrations
between 0.12 and 0.50 mg/kg. Eggs of three species
had concentrations above 0,5 mg/kg, namely a
butcherbird (1.34 mg/kg),.a brown tharnbill (0.69
mg/kg) and a superb fairy-wren (0.62 mg/kg). All
three species came from near cleared cultivated
areas,
Three species had eggs taken rom separate nests
at different times. In the black-faced cuckoo-shrike
from two widely separated areas, the Beltana ege
sample from the earlier brood had no pesticides
(0) Number of birds examined.
present whereas the Finnis sample of November had
a concentration of 0.50 mg/kg DDT plus meta-
bolites, Two eggs from different broods of the
superb fairy-wren had a five-fold difference in
concentration (0.12 to 0.62 mg/kg) of the pesticide,
On the other hand the concentrations of DDT plus
metabolites in broods of the iwo rainbow bee-eaters,
taken six weeks apart at Tailem Bend were virtually
identical (0.03 and 0.02 mg/kg respectively). Most
birds, whose broods were sampled, are
predominantly insect and arthropod eaters and only
a few cockatoos supplement their diet with seeds
while two species, the common bronvewiny and
little quail have seeds as their prime food.
In the bird broods sampled (Table 4) those eggs
with the nil or lowest concentrations of pesticides
came from nests in the lower rainfall areas.. Those
birds with the higher concentration of DDT plus
metabolites were from higher rainfall areas and
hence more closely settled districts where cropping
practices were more intensive.
In only two of the 26 foxes was the sex recorded
and no observations were given on the estimated
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ages of the foxes ic, juvenile or adult. The foxes
Were faken in the course of a tapeworm survey and
it is presumed they were mostly older animals and
hot pups.
HCB
All 26 samples of fox kidney fat showed traces
of HCB. Thirteen foxes showed the presence of
dieldrin in the kidney fat. The highest concentration
of 0.67 mg/kg was present in a fox from Mannum
district, the next highest was 0.27 ng/kg in a fox
from the Port Lincoln area (Table §). The remaining
11 foxes had concentrations ranging from 0.02 ta
0.16 mg/kg dieldrin. Six enly of the 26 foxes
cxamined showed the presence of pp'DDT or
metabolites, the highest concentration of 1.34
mg/ke was in the same fox from Mannum which
registered the highest concentration of dieldrin,
Discussion
The amounts of pp’ DDT used in the
Saddleworth-Riverton area were believed to be the
highest used anywhere in South Australia at that
time and survey results from that area were
considered likely lo show the highest residues from
74
DDT in this State, The annual application of about
1,§ kg/ha of pp'DDT to about three percent of the
area is not high compared with application rates
o! four to six times heavier on greater areas of
cotton and robacco planted elsewhere in Australia
(Australian Academy of Science 1972).
The data on pesticide residues in Australian fauna
is limited to small numbers of some of the more
common species present in different areas.
Comparison of residues is difficult because of
different tissues analysed,
Of the 38 native and four exotic bird species from
South Australia, only 12 native and one exotic
species Were common to the 63 species previously
surveyed in Victoria (Australian Academy of Science
1972). Table 6 shows these in descending order of
maximum concentrations of DDT plus metabolites
found in any tissues.
The highest concentration, 160 mg/kg was in the
body fat-of an Australian kestrel and was also the
highest level recorded in six other raptor species
from Victoria. Best (1973) reported up to 12.15
mg/kg wet weight in bulked samples of body fat
of fork tailed (black) kites, raptors from the NU.
The recording of 3.94 mg/kg im the Australian
TABLE 5, Concentrations of pesticides in fat af foxes — miscellaneous areas — 1973,
Pesticide (mg/kg wet weight) DDI plus
Date Town or district HCB DIBLDRIN pp DDT DDE DDD metabo-
received lites.
8-i-73 Por{ Lincoln 0.006 0.06 = * J »
131-73 Port Lincoln 0.007 0.06 7 * * *
13-1-73 Fort Lincoln 0,007 0.06 * } * *
25-4-73 Port Lincoln 0.08 0.14 * 0.08 ¥ 0.08
19-41-73 Port Lincoln 0.001 Q,27 * 0.02 + 0.02
25-v-73 Port Lincoln 0.02 . O11 0.26 * 0.37
28-v-73 Port Lincoln 0.18 4 > ‘3 »’ +
§-vi-73 Port Lincoln 0.2) + > O18 sf 0.18
16-vi-73 Murray Bridge 0.31 (1,05 > * * +
16-vi-73 Murray Bridge 0.006 a 4 ps *
16-vi-73 Mount P
Gambier 0.002 + 3 * *
25-¥i-73 Port Lincoln 0.008 0.08 + * * +
25-vi-73 Mout
Ganibie: 0.005 “ * ? * <
25-vi-73 Mount
Gambier 0.002 3 * * ‘ ?
27-vi-73 Murraylown 0.008 + * : 3 +
27-vi-73 Murraylown 0.015 0.04 bg 7 Le id
27-¥1-73 Murrayrown 0.014 (1.02 id . * *
2-vii-73 Port Lincoln 0.003 0.16 * 4 * 4
4-vii-73 Port Lincoln 0.004 * * * * 4
1B-vii-73 Mannum 0.028 0.06 * * . *
18-vii-73 Mannum 0.091 {).67 0,50 0.75 0.09 1.34
26-vii-749 Port Lincoln 0.004 * 4 + is &
26-vu-73 Port Lincoln 0.004 * * ® - if
26-vu-73 Port Lincoln 0.02 + ys 0.05 * 0.05
8-viii-73 Jamestown {).007 6.10 * 7 * ”
R-Vii-73 = Hallet 0,009 $ ‘ * « é
* Not detected
Th PR
RIR KS AoA Me OLSIN
Tan 6, Descending arder of concentrations of DDT plus ietabolites vi 13 bird spécres Conan to South Australia
and Victoria.
Species Maximunt pesnvide concentration mg/kg wel weight
South Australia Victoria
Australian kesrfel 3.94 160.0
Magpie lark TAKS v.20
Australian magpie 95.6 2.4
White faced heron At, 7 78.0
Line raven 0) 0.36
Laughing kookaburta 708 2)
Fairy penguin $2 0)
‘Stubble quuil 4.2 O.67
White-winged choigh a4 109
Great (larec black) cormorant (139 1%
Welcome switllow 0,80 1.2
Red wate bird 0,20 0.81
Starling ).60 0.07
kestrel is the only measurement From a rapror
species in S.A.
Although many waterbird species were examined
for pesticides in Victoria, only the white-faced
heron, the great (large black) cormorant and the
fairy penguin were also examined in the South
Aus|ralian studies and they had lower
coficentrations than their respective Victorian
SspECcIMeEns,
In the Namoi cotton growing area of N.SW,,
which had received the highest rates of application
ol pp'DDT in Australia (Australian Academy of
Science 1972), seven cormorants had ab average of
4.6.mg/kg (basis unspecified) in the body fat.. Fwo
freat cormorants from Moarook, '8.A., had 0.24
and 0.39 mg/kg wet weight in body fat whereas a
Victorian great (large black) cormorant had 1.8
me/ke wet weight in body fat.
Two quail from the Namo area bad an average
of 0.45 me/kg (basis unspecified) in body fat, three
stubble quail from Moonta,’S.A., bad 1.43, 1.47 and
4.2.me/kg wet weight in body fat and four stubble
quail in Victoria had 0,67 and 0.14 myg/ke wet
Weight in fat.
Two galahs (rom Saddleworth-Riverton area had
DDT residue concentrations of 4.7 and 6.0 me/kg
(stomach and liver fat respectively) whereas lower
concentrations (0,22 to 3.69 me/k wet weight),
were recorded in bulked samples of galahs fron
developed arid areas near Alice Springs and still
Tower concentrations (nil ta O17 mg/kg) in bulked
sainples from undeveloped arid areas in the N-T.
(Best 1973).
Freshwater crayfish from river and creek sources
in the Namoi area had DDT residue concentrations
ranging from 0.17 10 6.5 me/Ka, dry weight, and
higher vahies hetween 13,8 and 54,7 mg/kg,
averaging 29.9 mg/kg dry weight from specimens
collected in irtigation drains (Australian Academy
of Science 1972), A yabbie from a dam in the
Saddleworth-Riverton area had 4 Jower
concentrauion of 0:4 me/Kg wet weight..
In the South Australian results, pesticide levels
were generally higher in those bird species which
feed on Other animals, including insects, than in
bird species which utilise food of vegetable origin,
ie. Seeds, fruits, bulbs and other underground
storage organs,
The Jower proportion of pp'DDT than DDE
present in the tixsucs is indicative of pesticides
having been applied some time ago rather than of
recent application. There was no-apparent effect on
the general well-being of the birds examined in the
Saddleworth-Riverton survey although some, ag.
the sparrow, carried high concentrations of
pesticide, particularly DDE, Davis (1967, 1974) drew
attention to the wide variations i) susceptibility of
different bird species to specific pesticides and
considered that lethal levels for a species may vary
with conditions such as stress, From the high
concentrations of pp’DDT plus metabolites in the
butcher bird and its known contact with pp'DDT,
iL is presumed that this chemical caused its death.
With the sacred kingfisher from Rendeisham, the
cause of death is uncertain because there was Wo
known association with pesticide application and
it had lower concentrations of pesticides than three
other apparently unaflected bird species with high
pesticide concentrations in the Saddleworth-
Riverton survey (Table 1).
In the species, other than birds, examined,
concentrations of SDT plus metabolites were low-
In foxes the low concentrations. and absence (21 out
of 27) was interesting particularly in view of the
omiuvorous dict of (his species. Two foxes in the
Northern Territory also had low concentrations of
DDT residues (0.03 mp/ko) (Best 1973).
The DDT plus mctabolites-concentrations in eggs
PESTICIDES IN SOU'TH AUSTRALIAN FAUNA 77
of 23 South Australian bird species were relatively
low. Eggs of seven species were free of pesticides,
13 species had concentrations of 0.5 mg/kg wet
weight or less and the eggs of three species
contained 1.34 mg/kg (grey butcherbird), 0.69
mg/kg (brown thornbill) and 0.62 mg/kg (superb
fairy wren) (Table 4).
In the Victorian survey, eggs from nests of 14
different bird species, nine of which were water
birds, contained DDT plus metabolites. Four of the
species contained concentrations higher than values
found in eggs of the 23 South Australian species.
The eggs of a stilt had a concentration of 12 mg/kg
wet weight, a whistling eagle 3.2 mg/kg, a little pied
cormorant 2.5 mg/kg and a starling 1.70 mg/kg.
Because of wide differences in food preferences
and feeding habits of birds from Victoria and those
from S.A. there is no basis for comment except to
record the concentrations of DDT plus metabolites
found in the eggs of birds in these two surveys.
References
AUSTRALIAN ACADEMY OF SCIENCE (1972) The use of
DDT in Australia, Reports of the Australian Academy
of Science No. 14.
Best, S. M. (1973) Some organochlorine pesticide
residues in wildlife of the Northern Territory, Australia,
1970-71. Aust. J Biol Sci, 26, 1161-70.
Butcuer, A. D. (1965) Wildlife hazards from the use of
pesticides. Australas. J Pharm. 46, Suppl. 35, 105-9.
Davis, B. N. K. (1967) Recent developments in pesticide-
wildlife studies. J ent. Soe. Aust. (N.SMW.) 4, 3-9.
(1974) Advances in pesticide-wildlife studies since
1967. Ibid 8, 22-31.
Oxsen, P, & SETTLE, H. (1979) Pesticide contamination
of water rats in the Murrumbidgee Irrigation areas, New
South Wales, Australia, 1970-72. Pestic. Monit, J. 12(4),
185-188,
ECHINOCEPHA LUS OVERSTREETI DEARDORFF & KO, 1983
(NEMATODA:GNATHOSTOMATOIDEA) FROM ELASMOBRANCHS AND
MOLLUSCS IN SOUTH AUSTRALIA
BY IAN BEVERIDGE
Summary
Specimens of Echinocephalus overstreeti were found in the following elasmobranch and
chaemeriform fishes from South Australia: Heterodontus portusjacksoni (Meyer, 1793),
Parascyllium ferrugineum McCulloch, 1911, Orectolobus maculatus (Bonnaterre, 1788),
Myliobatis australis Macleay, 1881, Aptychotrema vincentiana (Haacke, 1885), Trygonorhina
guanerius Whitley, 1932, Raja whitleyi Iredale, 1938, Urolophus mucosus Whitley, 1939, Dasyatis
brevicaudatus (Hutton, 1875), D. thetidis (Waite, 1899), Hypnos rnonopterygium (Shaw & Nodder,
1795) and Callorhynchus milii Bory de St Vincent, 1823. Gravid nematodes were found only in H.
portusjacksoni. A redescription of the nematode is given and its differentiation from congeners
discussed. Nematodes undergoing the final moult in elasmobranchs permitted the identification of
the scallops Pecten albus Tate, 1887 and Chlarnys bijrons (Lamark, 1819) as possible intermediate
hosts.
KEY WORDS: Nematoda, Gnathostomatoidea, Echinocephalus, morphology, life history.
ECHINOCEPNALUS OVERSTREETI DEARDORFF & KO, 1983 (NEMATODA:
GNATHOSTOMATOIDE A) FROM ELASMOBRANCHS AND MOLTEUSCS IN
SOLTH AUSTRALIA
by TAN BEVERIDGE*
Summary
Hevreipoe, 1. 987) Evhinocephalus oversireeti Deardortt & Ko, 1983 (Nematoda! Gnathostomatoidea)
from clasmobranchs and maltuyes in'South Australia. Trams, 8, Soc. & Aisi. W112), 79-92, 29 May, 1987,
Specimens of Echinoeephalus overstreett were found in the following elasmobranch and chacmeritorm
fishes fiom: South Australia: Hereraduntus portusjackson’ (Meyer, 1793), Paraseviliam ferruginewn
McCulloch, 1911, Orectolobus maculatus (Bonnaterre, 1788), Mylobatis australis Macleay, 188),
Apiehotreme vincentiand (Haacke, 1885), Trygorortina guunerius Whitley, 1932, Raja whirley) ledale,
1938, Urolophus tnucosus Whitley, 1939, Dasvatis brevicaudatiuy (Hution, 1875), D. thetidis (Waite, 1899),
Hypnos monoplerygine (Shaw & Nodder, 1795) and Callorkynchus milii Bory de. St Vincent, 1823, Gravid
nematodes were found only in J, portusjacksori, A sedeseription of the nematode is given and its differen-
clailan from congeners discussed, Nemarodes undergoing the tinal moule in clasmobranchs permirted the
identiligation of the scallops Pecten albus Tae, 1887 and Chtanis bifrons (Lamatk, 1819) as possible
intermediate hosts.
Kry Worns, Nematoda, Ginathostomarolidea, Kehinoeephalus, morpholagy, lite history,
Introduction
Adults of the nematode genus Echinovephalys
Molin, 1858 occur in the stomach and spiral valve
al sharks and rays in various regions of the world,
Larval stages occur commonly in the gonads or
adductor muscles of molluscs as well as it the
mesentenes of teleosts, In Austrahan waters, adult
Echinocephalus Wave been found in the Port
Jackson shark, Heleradonius portusjacksoni
(Meyer 1793) (syn. ¢7, philipp/) and the stingaree,
Urolaphus testaceus (Mueller & Henley 1841)
Clohnston & Mawson 1943, 1945a), Johnston &
Mawson (1943) assigned the material they examined
lo £ spinosissimus (Linstow 1904), a species
described originally from Myliohalis aquila
(Linnaeus 1758) from the Gulf of Manaar, Sri
Lank#. However, £ spinosissimus has usually been
regarded asa synonym of E, uneinarus Molin, (858
(see Milleman 1963). Johnston & Mawson (1943)
noted however thar their specimens were much
Jarger than any previously described, while
Milleman (1963) pointed to differences in cephatic
anatomy and the caudal papillae of the male tail,
suggesting that the Australian specimens might
represent a distinc! species, Notwithstanding this
suggestion, Milleman (1963) tenlatively inched
Johnston and Mawson’s specimens under &.
dutemnals,
Larval stages of Eehinocephalus in the Australian
rewon have been found in the molluses A riviera
bulfoti Bernardi, 1861, Aatylesia scalarine (Lamarck,
(S18) and Polinices conicus (lamarck, 1822)
Unhnston & Mawson '945ar Lester #/ a/, 1980) as
well as in che mesenteries of the teleosts
Chrysophrys auratus (Schneider, 1801) (Syn,
Pagrosomus auratus), Platycephalas arenarius
Ramsay & Ogilhy, (885, A bassensis Cuvier, 1829,
P fusens Cuvier & Valenciennes, 1829, P laevigarus
Cuvier, 1829, Psendolabrus psittaculus (Richardson,
1840) and Sillaginndes punctatus (Cuvier, 1829)
(Johnston & Mawson 19484, b; Hooper 1983),
Larval echinacophalids have.alyo been found in the
stomachs of turtles, Carefta caretia (Linnacus, 1758)
(Lester et al 19R0), and dolphins, Delphinas
delphius Linnaeus, 1758 (Johnston & Mawson
1941). Generally, specitic identifications of larvae
have not been made excep! for those identified as
E. uncinatus by Johnston & Mawson. (1945a, b)
based on the earlier designation by Baylis & Lane
(1920) of a larval specimen as the type of Molin’s
species uncinatus. Baylis & Lane's (1920) action is
now considered invalid (Milleman 1963; Beverigc
1985) but in the intervening period many larval
forms have been incorrectly identified as £,
uncerains (see Beveridge 1985).
The taxonomic status of both larval and adult
forms of Bchinocephalus from Australian hosts is
therefore i duubt, Examination of new collections
has shown that they belong neither to £, wacinaties
nor to £. spinasissimus, but to &. oversrreet/
Deardorff & Ko, 1983 originally described from the
tay Ja@eriura sielanpspilos Bleeker, 1853 from the
Marqueses Islands (Deardorff & Ko 1983) although
they differ from the original description in some
* Central Veterinary Laboratories, South Austrahan
Departinent of Agriculure, ove Instiruw of Mecical
and Veverinary Scienve, Frome Road, Adelaide. S, Aust
5000.
a IAN BEVERIDGE
jeapects, A detailed description of both adult and
lurval specimens from Australia is [herelore given
below, tegether with some observaliens on life
Iistony..
Materials and Methods
Elasmebranchs were collected on cominercial
fishing vessels in St Vincent and Spencer Gufs,
Encounter Bay, off Beachport and off the south
coas!| of Kangaroo Isfand, S.A. Additional
specimens were obtained af Goolwa, 5,A,, using
tramllines, The sex and teral lenerh of each fish were
revorded and the gastroinicstinal iraet was removed
and opened. Nematodes were presetved either by
washing Them in sea-waler and fixing them
immediuiely in Rerland’s Wnid or allernalively,
opened spiral valves and stomachs wete flooded
with boiling waler Lo straighten any helminths, and
formalin was added to preserve the specimens.
Nematodes were cleared in lactophenol and the total
length measured while cleared, Drawings were made
using a drawing tube attached to an Olympus BH
microscope. Measurements were made cither
Girectly with an ocular micrometer ar trom cali-
brated drawings: In the descriptions, measurements
ure given in millimetres as the range tor Tive
specimens followed by the mean in parentheses,
Scallops, Chlamys (Equichlamps) difrass
fLumarck, |809) and Pecten albus Tate, 1887
(probably a synonyin of PB meridionalis Tate, 1887),
were collected from Nurthayen and hdijhburgh. The
otal length from hinge te free side, was recorded
and they were then examined fresh of were frozen
prior to examination jor nematodes. Live larvae
were fixed in Berland’s fluid and cleared in
lactophenol, Larvae from frozen scallopy were fixed
in 10% formol saline. Sixteen abalone, Hefioris
Jeevigate Donovan, 1808 frum Edithburgh were also
examined.
Larval and adult nematodes were prepared for
seaming electron microscopy (SEM) by dehydiating
in ethanol, air drying and couting with gould an¢l
curbon. Specimens were viewed with a JEOLSE
microscope.
All nematodes collected have been deposited in
the Australian Helininthological Collection (AHC)
heused in the South Ausinalian Museu.
Existing collections of Evkinucephalus in AHC
were alsu examined as well as the paratypes of E.
oversireeti trom the United Stales National Muscum
Helhuinth Collection (77384),
Adult fermale nematodes {rom one shark were
inaintained in sca-water at room temperature For
the collection of caps. Eag development and
hatching was observed. Larvae were lined in fori
saline or in lactaphenal.
Results
Echinocephalus averstreeti Deardort? & Ko, 1983
FIGS 1-38
Adults: Large sroawt nematodes; body woarmed,
maximum wideh in mid-body region; cuticle with
fine, regwlarly-spaced annular striations 0.002-0,003
apart; female straight; tail of male coiled in loose
spiral. Mouth opening derso-ventrally elongate with
2 elongate tateral pseudolabia;y lateral part of
pscudolabium bulbous, with median amphid and
double cephalic papilla on enter side; medial
exuemity of pseudolabium wider than lateral
extremity, indobed. cach lobe with 2 cuticular
thickenings along external edges; thickenings of
median lobes interlock 10 give appearance of pairs
of “teeih” in amecal views median lobes may be
partially or fully introverted such that one lobe or
both is wot visible in apical views projections or
“igeth™ of sub-dorsal and sub-ventral lobes of
pseudolabia only clearly visible in mediaiy views,
sumefintes visible in oblique or apical views giving
appearance of additional interlocking ‘Yeeth”;
postero-dersal and postero-ventral part of buxe of
each pseudolabium with distinet cuticular serra-
tions; small triangular interlabia present dorsally
ind ventrally, with 2 additional small (riangular
projections on cither side of apex, Cephalic bulb
prominent, anned with numerous cows ol sntall
uncinadte spines; anterior rows incomplete, restricted
to dorsal and ventral areas of bulb; rows af mid-
bulh region frequently non-conlinuous; raws of
spines won-overlappings buccal capsule weakly
developed; oesophagus divided into antenur
muscular und posterior glandular sections; nerve
ring in anterior oesophageal region, immediately
posterior to cephalic bulb; 4 cervical sacs extend
tram cephalic hulb almost to junction of muscular
and glandwar ocsophagus. Tail conical, with blunt
fips tail (ip without ernaieitatian,
Male: Votal length 53-60 (50); maximum width
1,0-1,2 (I. cephalic bulb 0.31 0.56 (0,54) long by
0.76-0.82 (0.78) wide; bulb with 30-40 (34) rows of
spies; spires 0.010 long; oesophagus 5.9-8,5 (7.2)
long; anleriur muscular region 3.0-5.4 (4.7) long:
eee EEE
Pips 1-10 Achieavephafus aversteert, 1. Oesophageal region, lareral wew; 2. Cephalic extremity, lateral views 7
Pseudolabium, lateral view; 4. Pscudolabium internal suiface; 5. Pseudolabia, apical view; 6. Pseudotabia, dorsal
view; 7. Spines of cephalic bulb; 8. Dettils of interlocking tveth of pseudoktbia, apicd view; 9 Dvirid, median
view) TD, Transverse section through oesophageal region showing (yar cervical saves. Scule Limes in cum.
31
E. OVERSTREET] IN VISH AND MOLLUSCS
1h 7
vl
i} Wit.
wee
82 IAN BEVERIDGE
we)
se Sa
SHO
Pt
wks
4 11,12,18 \
o. 45.16
ey |
& OVERSTREET? IN FISH AND MOLLUSCS &3
posterior plandular region 2.5-3.2 (2.9) long; nerve
ting 0.85-0.9§5 (0.91) from anterior end: dcirids
1.1-1 3 (1.2) from anterior end; cervical sacs extend
2.6-3.2 (2.9) posterior [0 cephahe bulb, Tail }.0-1.6
(1.3) long; spicules sub-equal 1,70-1,94 (1.82), ratio
Of lett: right spicule lengths (.98-1.03; distal
extremity of spicule slender, tapering to sharp point;
gubernaculum present, poorly sclerotised, clearly
visible in young specimens only, punctate, 0.07-0,09
(0.08) long; W shaped in ventral view. Prominent
caudal alae present on male tail, extending 1.6-2.6
(2.0) from posterior end; prominent rugose area
extending along lateral areas of alae fram between
Papillac 2 (numbered antero-posteriorty) arid 3 to
between papillag § and 6, most prominent in region
of papilla $; composed of prominent! bosses
arranged in irregular rows, wiving cobblestone
appearance, 7 pairs of caudal papillae; 3 pairs pre-
anal; anterior pair distant from remainder; pairs
2 and 3 on lateral alae, pait 2 largest; 4 pairs of pust-
anal papillac; pair 5 slightly medial to remainder;
pairs 6 and 7 witlely separated from pair 5; pair 6
larger than pair 7, phasmids ar level of pairs 7,
Annularions on ventral surface of male tail; extend
from a little anterior to papillae to between mugose
areas, Annulations 0.007-0.010 (0.009) apart; meet
normal striations in region of lateral lines.
Jreriaiiere mates: Total length 20-31 (27); maximum
width 0.53-0.68 (0.60); cephalic bulb 0.39-0.42
(0.40) lung by 0.56-0.64 (0.60) wide; bulb with
30-35 (32) rows of spines: spines 0.010 tong:
oesophagus 4.2-5,.5 (4.7) kong; anterior muscular
region 2.8-3.5 (3.0) long; posterior glandular region
1,2-2.0 (1.6) lung; nerve cing 0.55-0,74 (0,64) From
anletior end; deirids 0.77-0.90 (0.84) from anterlot
end; caudal alae extending 1.1-1.5 (1.4) from
postenor end,
Female; Total length 58-63 (39); maximum width
1.3-1.6 (1.5); cephalic bulb 0.48-0,66 (0,55) long by
0,80-0,90 (0,84) wide; bulb with 29-35 {33} rows of
spines; spines O.0KS-0.010 (0.00%) long; vesophagus
8,0-8.9 (8.5) long; anterior muscular region 5.0-6,0
(5.5) long; postertor glandular region 2.4-3,3 (3.0)
long; nerve ring 0,70-0,35 (0.77) from anterior end;
deirids 0,88-),20 (0,96) from anterior end; cervical
sacs extend 2.1-3.8 (3.1) posterior to cephalic bulb;
tail 14-17 (1.6) long, conical, blunt; vulva 1.8-2.6
(2.3) from posterior end; vagina (sphincter to vil vis)
0.9-1.2. (1.0) Long; length from sphincter to division
of iterus approximately 6.0; uterus didelphic,
prodeiphic; ¢gks oval 0.050-6,055 (0.053)
«0.036.040 (0.038), Operculate, with ifregularly
pitted shell.
hnmeure feioles Total length 28-31 (29);
maximud width 0,58-0,86 (0.66); cephalic bulb
0,34-0,45 (9.39) long by 0.56-0,72 (0.64) wide; bulb
with 30-345 (33) rows of spines 6.010 long;
oesophagus 4.2-6.5 (5,1) long; anterior muscular
region 2.8-4,8 (3.6; posterior glandular region
1.4-2.7 (17) nerve ring 0.67-0.72 (0,70) from
anterior end; deinds 0,79-1,02 (0,89) from anterior
end; cervical sacs cxtend |.9-2.9 (2.3) posterior tu
cephalic bulb. Tuil 0.95-1.06 (0.98) long; vulva
[,35-1.61 (1.48) from posterior end} vagina
(sphincter to vulva} 0.51-0.60 (0,56); eggs absent,
Fourth-stage farvee: (measurements of specimens
from Chlamps bifrens. Small nematodes, coiled
ventrally; 12,9-14,3 (13.7) long, maximum width
0,28-0.35 (0,33); body covered with fine annular
striations; mouth opening dorso-ventrally elongate,
sutrounded by 2 lateral pscudolabia; pseudolabia
without accessory lobes, beating amphids and pairs
of barely discernible cephalic papillac; cephalic bulb
prominent, 0.25-0.40 (0.33) long by 0.30-0.36 (0,33)
wide; 5 tiny spine-like structures, arranged in rows
of Zand 3 on dorsal and ventral aspects of jnouth
opening. Cephalic bulb armed whh 6 rows of
subulate hooks arranged in semi-cireles, with
unarmed spaces laterally: hooks yenerally increase
in size medially, but some median hooks smaller
than submiedians; number of hooks in dorsal and
ventral sectors of rows not always identical; hook
numbers as follows: row 1, 14-18 (16,9); row 2, 16-20
(17.4); row 3, 16-22 (18.8) Tow 4, L6-£9 (18.3):
tow 5, 16-19 (18.2); hooks tn tow 6 could not be
counted accurately, Length of median hooks
Increases from row | to row 4; jnedian hooks of
rows 4-6 of similar size; hook lengths: row L,
0.020-0,025 (0,023); row 2, 0.025-0.035 (0.031); row
3, 0,020-0,035 (0.034); row 4, 0.035-0.040 (0,038)
row §, 0.035-0.045 (0.039); row 6, 0,035-0,040
(0,038; oesophagus 2,2-4.4 (3,3) long: tail conical,
0,25-0,36 (0.30) long. Genital primordia visible in
some specimens. Male primordivm divided into 3
regions; posterior region terminating near secrum,
contains approximately § cells; mit-regtan short,
with 6 pairs of cells; anterior region elongate, cells
atranged in pairs tn posterior two-thirds, singly in
anterior third, Female primordium elongate,
subdivided at anterior cnd into two short lobes, each
with 5 cells.
Aegs! Eggs held in seawater at room teniperature
began te hatch after 10 days. Larvae emerging from
bigs H-1. Achthacephalus overstreesi, 11, Male cail, ventral view; 12. Male tail and spicule, lateral view; 13. Detail
of nigose area bn vicinity of Jeft fourth caudal papilla of mole; 14. Spictile tip, lateral view, 18. Gubermaculue,
lateral view: 16, Gubernaculum, spicule sheaths and spicule tips, ventral View; 17- Ternvinal region of female genitalia,
lateral view, IR_ Female tail, lateral views 19. Ege. Scale ines In mien,
84 IAN BEVERIDGE
wi 5 OTD,
Fagg 5 9) dee
ne | asta
94 a8
ay 48
Qe Gam
21,23,25,26
Figs 20-26, Larva of Fchinacephalus overstreeti trom scallops, Chiamys bifrons, 20, Cephalic extremity, lateral view;
21. Median row of hooks, lateral view; 22. Ocsophageal region, lateral view, 23. Mouth opening and pseudolabia,
apical view; 24, Tail of female, lateral view; 25, Pseudolabia, dorsal view; 26. Pseudolabia, lateral view. Scale lines
in mm.
E. OVERSTREET! \N FISH AND MOLLUSCS 85
Figs 27-29. Larval stages of Echinocephalus uncinatus. 27. Male genital primordium in larval stages from scallop;
28, Female genital primordium in larval stage from scallop; 29, Ensheathed larva newly emerged from egg. Scale
lines 0.1 mm.
eggs were ensheathed, with a conspicuous
thickening at the anterior end of the sheath. Few
internal features of the larvae were discernible
except for an apical thickening and paired granular
regions in the mid-region of the nematode body,
even when stained with methylene blue. The tail of
the larva was long and slender with a recurved tip.
Motile larvae quickly became attached, by the tail,
to debris in the sea-water. Length of larvae
0.108-0.122 (0.114), maximum width 0.009-0.010
(0.009).
Fourth-stage larvae in elasmobranchs: Larvae
identical to those occurring in scallops were found
in the stomachs of rays and sharks, In addition,
several specimens were found undergoing the final
moult. The external cuticle of the cephalic bulb bore
six rows of hooks identical with those occurring on
larvae in scallops; underneath the external cuticle
were the numerous rows of small spines found on
the cephalic bulb of the adult, indicating that the
larvae in scallops and the adults in elasmobranchs
belong to the same species. No further moults were
observed in elasmobranchs.
Spicule lengths: Observations indicated that the
spicules of immature nematodes were shorter than
those of adults. Data (Fig. 39) indicate a linear
relationship between worm length and spicule
length of the form y=0.46+0.24 x (r? =0.922).
Larval stages in scallops; Larval stages were found
in C. bifrons and P. albus from Northaven in Gulf
86 IAN BEVERIDGE
34
Figs 30-36. Micrographs of adult and larval Echinocephalus overstreeti. 30. Cephalic end of adult with single interlabium
exerted; 31. Cephalic spines of adult showing discontinuities in rows; 32. Cephalic end of adult with both interlabia
visible; 33. Cephalic end of adult with both interlabia inverted; 34. Cephalic end of larval stage from scallop, showing
detail of interlabia and groups of spine-like structures anterior to major rows of hooks; 35. Larval E. overstreeti
newly emerged from egg; 36. Egg shell showing mammillate surface. Scale lines: figs 30, 32, 33, 34, 0.1 mm; figs
31, 35, 36, 0.01 mm.
St Vincent and in C. bifrons from Edithburgh The prevalence of infection was significantly
(Table 1). A higher prevalence of F. overstreetiwas lower, using a x? test, in C. bifrons less than
found in P albus as well as a higher intensity of 65 mm in total length compared with larger class
infection. sizes. Other differences in prevalence between class
E. OVERSTREETI IN FISH AND MOLLUSCS 87
Figs 37-38. Adult male and larva of Echinocephalus overstreeti. 37. Male tail, ventral view with papillae numbered;
phasmid indicated by arrow; 38. Cephalic end of larva from scallop, Ch/amys bifrons. Scale lines 0.1 mm.
2,0
Y= 0,46 +0.024x
mm
length
1.0
Spicule
12) 10 20 30 40 50 60
Nematode length mm
Fig. 39. Relationship between spicule length and total body length of 40 specimens of Echinocephalus overstreeti
from Heterodontus portusjacksoni.
88 IAN BEVERIDGE
TABLE 1. Occurrence of larvae of Echinocephalus overstreeti in scallops, Pecten albus and Chlamys bifrons.
Intensity of
Prevalence infection
Host species Locality No, examined (%) Range (mean)
Chlamys bifrons Northaven 136 61.8 1- 7 (2.0)
Edithburgh 34 41.2 1- 4 (2.5)
Pecten albus Northaven 20 80.0 1-14 (4.4)
ee
sizes were not significant statistically. Intensity of
infection (Table 2) declined slightly, but not
significantly, with increasing size.
The occurrence of different intensities of
infection fitted an exponential curve of the form
y=113.6, -0.566 x(Fig. 40).
No nematode larvae were found in abalone
(Haliotis laevigata).
%
Prevalence
No.
larvae per scallop
Fig. 40. Intensity of infection in 153 scallops, Chlamys
bifrons, from St Vincents Gulf, with the larva of
Echinocephalus overstreeti, Numbers of scallops in each
class is indicated above the histogram. The relationship
between Prevalence and larvae per scallop is
y=113.6, 966%,
Definitive hosts: E. overstreeti, including some
gravid females, was found commonly in H.
portusjacksoni (Table 3). There was no association
between host length, used as an indicator of age,
and either prevalence or intensity of infection
(Table 3). Infections were also found in several
species of ray and in the chaemeriform C. milii, but
in none of these hosts were gravid females seen. The
largest nematodes found in a ray were in Myliobatis
australis in which females reached 45 mm in length;
none was gravid. No E£. overstreeti were found in
11 Furgaleus ventralis (Whitley, 1943), 11
Notorhynchus cepedianus (Péron, 1807), 40
Mustelus antarcticus (Guenther, 1870), 6
Galeorhinus australis (Macleay, 1881), 9
Pristiophorus cirratus (Latham, 1794), 15
Centrophorus moluccensis Bleeker, 1860, 41
Squalus megalops (Macleay, 1881), 17 Carcharhinus
brachyurus (Guenther, 1870), 30 Orectolobus
tentaculatus (Peters, 1864), 14 Narcine tasmaniensis
Richardson, 1840, 47 Raja cerva Whitley, 1939, 20
Urolophus expansus McCulloch, 1916, 8 U.
cruciatus (Lacépede, 1804) and 7 Squatina australis
Regan, 1906 examined. Histograms of nematode
lengths (Fig. 41) indicate that those from rays were
predominantly immature nematodes and that a
greater proportion of the nematodes recovered from
rays were larvae.
Echinocephalus from other regions of Australia:
Two collections of Echinocephalus from
Queensland were available for examination. The
first, consisting of 15 males, 25 nongravid females
and 2 larvae of E. overstreeti was collected from
Dasyatis fluviorum Ogilby, 1928 at Caloundra
(AHC 1077).
A second collection consisting of 5 females and
2 males, taken from Aetobatus narinari (Euphrasen
TABLE 2. Prevalence and intensity of infection of Echinocephalus overstreeti /arvae in scallops, Chlamys bifrons
of different length classes.
Scallop length Prevalence Intensity of infection
class No. of infection (no. larvae per infected
(mm) examined (%) scallop)
<60 20 40 2.88
61-65 23 48 2.18
66-70 31 71 2.18
71-75 52 65 2.03
>76 27 63 1.94
eC rrr
&, OVERSTREETI IN FISH AND MOLI USCS 89
Taste 3. Prevalence and intensity of infection af adult Echinocephalus overstreeti #7 Heterodantus portusjacksoni.
Size class
Intensity of
Prevalence infection
{cm) No. sharks (fy) range (mean)
Males 30-40 13 100 8-52 (22)
41-S0 y 100 2-61 (19)
51-60 5 BO 3-13 (7)
Gls 3 77 12-16 (14)
Total 30
Females 30-40 10 100 §-35 (19)
4t. 50 9 89 3 47 (14)
51-60 5 100 &-30 (13)
61= 5 40 13-63 (38)
Total 29
TABLE 4. Occurrence of Echinocephalus overstreeti in elasmobranch and chaemeriform fishes from South Australian
gulf and adjacent waters.
Tatensity of Adult (A)
Prevalence infection or larval (L)
Host species No. Examined (%o) Range (mean) stages
Heteradontius portusjacksoni 49 ROS 3-63 (13.9) AL
Parascyllium ferrugineurn 6 66,7 1-13 (8,3) {1
Oreclolobus maculatus 7 14,3 2 L
Dasyatis brevicqudatus 17 35.3 1-42 (8.3) L
Dusyatiy thetidis 4 75.0 1-10 (2.7) L
Raja whitleyi 10 40.0 1-24 (3.1) AL
Urolophus mucosus 10 20.0 1-4 = (2.5) AL
Trygonorhina guanerius 47 12.7 1-20 (7.7) AL
Aptychotrerna yincentiana 35 2.9 1 L
Myliobatis australis 26 30.8 1-16 (3,6) AL
Flypnos monoprerygium 6 16.7 1 L
Callarhynchus milti 21 42.8 1-4 (2.0) L
1790) from Moreton Bay (AHC was
identificd as £. sinensis Ko, 1975.
1080),
Discussion
The adult nematodes described above mast
closely resemble &. overstreeti, They are distin-
guished from all congeners except E. pseudounci-
natus Milleman, 1951 by the arrangement of the
papillae on the tail of the male, with the terminal
three pairs widely separated from the remaining
papillac. In addition, the specimens differ from E.
diazi Troucy, 1969, EL multidentatus Balyis & Lane,
1920, E. southweili Baylis & Lane, 1920, E. pseucto-
uneindlus and E, sinensis Ko, 1975 in having more
than 29 rows of spines on the cephalic bulb, from
E. spinosissimus (Linstow, 1905) and £. daileyi
Deardoff, Brooks & Thorson, 1981 in possessing
a well developed rugose area on the ventral surface
of the tail of the male and from &. uncinatus in
the form of the rugose area, which in the latter
species is ornamented with regular tows of fine
bosses (Beveridge 1985) rather than the irregularly
shaped bosses illustrated in Fig. 13. BE. mobulae
Kalyankar, 197] is considered a species inquirenda
following Ko (1975) and Soota (1983), as is &.
aligocanthus Anya, 1977 which is based solely on
a Jarval form, These two species have nor been
considered further. The Austrahan specimens
conform to. the description of E. averstreeti in all
details except for measurements, Males. from the
type series of &. overstreeti measured 21-30 mm
with spicules 0.4-1.4 mm long (Deardoff & Ko
1983), compared with Australian specimens
measuring 55-60 mm and. spicules 1.70-1.94 mm.
However, as indicated in Pig. 39, considerable
growth occurs during the final stages of
development and the spicules as well as the body.
increase markedly in length, the relationship
between the two being linear. The ratio of spicule
length: body length for the Australian material is
similar to that of &. overstreeti. Specimens of E.
overstreeti from Australian rays were generally
smaller than those from sharks, and since the type
series of E. overstreeti was taken from a tay,
90) 1AN BEVERIDGE
LARVAE MALES
No nematodes
5 to % 5S 10 1 20 25 39 435 4y
Size
45 50 55 60 65 70 s
FEMALES
H.goriusjacksani
Other elasmobranchs
and chaemeriforms
w 1S 20 25 3O 3s 40 45 50 Ss €6 €5 7H
class mm
Pig. 41. Size distribution of larval and adult Behinocephalus averstreet] arranged in § mm size classes, Numerals
on abscissa indicate lower limit of size class. Nematodes from Heterodontus portusjacksoni are compared with
thase from other species of shark, from rays and from chaemeriforms, grouped together.
Taeniura melanospilos, the size differences noted
may be host induced. The Australian material was
compared with paratypes of E. overstreeti before
being, assigned to this species.
Redescription of the cephalic anatomy of
Australian specimens confirms that the
arrangement of the pseudolabia and teeth is
identical with congeners and contradicts
Milleman’s (1963) suggestion that the pseudolabia
and tecth of Australian Aehinecephalus are unique.
These differences plus the large size of the
specimens was the basis of Milleman’s suggestion
that Johnston & Mawson's (1943) Specimens
tépresented a new Species. Although a correct
conclusion, Milleman’s (1963) reasons for reaching
it were not.
Two features of the morphology of the adult, the
arrangement of the pseudolabia and the guber-
haculum, warrant comment. Examination of several
specimens of E, overstreet? by SEM revealed that
cephalic morphology is cxtremely plastic, a feature
nol noted previously m the genus. The median lobes
of the pseudolabia may both be exerted and inter-
lock (Fig. 32), one may be concealed below the
other (Fig. 30), or both may be completely intro-
verted such that the trilobed nature of the inter-
labia is concealed (Fig. 33). Congeners have been
described exclusively with both pseudolabia exerted,
The gubernaculum was only clearly visible in
immature males, as a weakly sclerotised structure
at the junction of the spicule sheaths, In adult
males, the highly developed surrounding muscu-
Jature makes it more difficult to detect, A
gubernaculum is present in all species which have
been described or re-cxamined in. recent. years
(Troncy 1969; Ko 1975; Deurdorff et af 1981;
Deardorff & Ko 1983; Beveridge £985), and may
well be present in all members. of the genus.
Attempts to sub-divide the genus based on the
Presence or absence of a gubernaculum (Deardor!f
et al, 1981) may thus be premature until all species
have been re-examined carefully,
The larvae of Echinocephalus from scallops are
also attributed to E. overstreeti based on the
evidence of moulting nematodes which clearhy
demonstrated the features of the fourth silage
externally and those of the adult internally,
Milleman (1963) considered the stage of Ff.
pseudouncinatys in abalone to be the second stage,
and Ko (1975) described the larval stages of E
Sinensis in oysters as second and third stages. By
contrast, the nematodes in scallops are considered
to be fourth stage larvae. Not only are adult features
present in larvae entering the final moult, but well
developed genital primordia are present in larvae
from the scallops. Gonad development normally
E, OVERSTREET IN FISH AND MOLLUSCS 9)
occurs during the latter part of the fourth stage in
teinarodes. Although the mematodes jn scallops
undergo considerable growth in H. portusjavksoni,
no evidence of moulting was seen in nematodes
greater than 15 min in length when the six rows of
hooks on Lhe cephalic bulb are replaced by the 30
rows present in the adult.
Only the larvae of £. psevdouneinatus and £
sinensis have been described tn deiail, and both are
remarkably similar to that of E overstreeti. Hook
numbers and sizes may allow rhe species ta be
separated, bul more detailed niorphological data
are needed before this can be done, No larval
Echinecephalus were found in the abalone
examined, |huugh abalone are the normal pnter-
mediate host of & Pseudauncinatus (Milleman
1931), E. overstreert is the only species that has been
found in South Australia, and on this provisional
basis, earlier records of Echinecephalus sp_ or E.
wnermanes from this region can probably be
attributed to E. overstreeti. Records from other
parts of Australia (Johnston & Mawson 194Sa, b;
Lester ef al. 1980; Houper 1983) cannot be assigned
to any species with certainty at present, pending
investigations of the geographic distribution of &.
overstreeti, but the original description of the
nematode from the Marquesas Ishands sugvests that
iLis widely distributed. The naming of larval forms
as &, uncinatus by Johnston & Mawson (1945a, b)
is due to the fact chat at che time of their work,
the name sencinatus had been applied, incorrectly
by Baylis & Lane (1920), to a larval nematode from
Dasyatis centroura (Mitchill, 1815) (syn. Tygon
firuceo) from the Adrialic sea (Milleman 1963;
Beveridge 1985) and their specimens closely
resembled larval wacinerus &. uncinarus is known
only from the Adriatic and Black seas (Heveridge
1983).
Ko (1975) described the larva of & sinensis as
having seven rows of spines, &, overs(reeti ts
considered to have six major rows, and the groups
of five spine-like projections dorsally and ventrally,
posterior to each pseudolabium, described as the
“first" row by Ko (1975) are not considered to
constitule a ruw of spines, Their siructure ts quile
different to spines of the major rows, but their
function is not known.
Data presented above suggest that Ff, porius-
yweksont is the principal definitive host of &.
oversireeti in South Australian gulf waters. This
shark feeds predominantly on molluscs, echino-
derms and crustaceans (McLaughlin & O'Gower
{97]), and gravid female nematodes were commonly
encounrered in it. There appenred to be a slight
reduction in prevalence in larger sharks of both
sexes, Hithouoh the number of sharks examined in
cach size class was small. A. pertusjacksonl are
Approx, 23 em at birth, and grow at 5.1-7.4cm per
year. (McLaughlin & O°Gower 1971), Thus mose of
the small sharks examined were probably at lease
Two years of age, and the relatively large numbers
of nematodes in sone sharks of this length ts
therefore not surprising. Because of the paucity at
data on growth in A. portusjeckson! (McLaughlin
& CyGower 1971), the relationships between host
age and infections with &. oversirevti has not been
analysed in detail. Scallops are heavily infected with
E. aversrreeti, but ne other species of moltuses other
than abalone Were examined, and scallops mav not
be the only passible intermediate hosts,
Rays and other species of shark which feed on
shellfish may also ingest larvae ol £) oversirées.
The high percentage of larvae present and the lack
of mature nematodes in hosts other than H.
portusjacksoni suggests that they may not be
entirely suitable hosts for the development al’ the
nematode. The largest nematodes found in a ray
were ina specimen of Myliabasis australis, bit the
females were not gravid. Examination of a wider
variety of elasmobranchs may reveal thar £.
oversireeti develops lo inaluriry in some species,
since the type specimens from the ray Teeniuee
malanospilos were gravid (Deardorff & Ko 1983),
The same species of ray occurs In northern
Australian waters but none were examined in (his
study.
E. sinensis ts reported here for jhe first time Fran
Austraha, in the ray Aetoduues nurinari. The
nematode was formerly known only Irom Hong
Kong, from the same definitive host species (given
as A. flagellum, now a synonym of A. arinart}.
The usual intermediate hosts are oysters,
Crassostrea gigas {Thunberg, 1793) (Ko 1975),
Acknowledgments
Thanks are due 10 Mesdames E. Moore and J.
Clarke and Messrs M, O'Callaghan and R. Martin
for experi assislance in the laboratory and for
collecting sharks and rays, to Mr B, Robertson for
the collection and <xamination of many of the
clasmobranchs and tu Dr J. R. Lachtentels Por tle
loan of specimens, MrK, Smith took the scanning
electron micrographs and Dr T, Deardorff kindly
conficmed the identity of the species.
The descriptive work was carried oul in the
Laboratoire des Vers, Muséum national d’Histoire
naturelle, Paris. Prof. A. G. Chabaud ‘s thanked
for providing laboratory assistance and financjal
support, Mr J. Glover is thanked for his advice on
host nomenclature and Dr D. M, Spratt for
comunents on a draft of the manuscript.
Collection of the majority of rhe elasmobranchs
was supported financially by tbe Austraiisn
Biological Resources Survey,
92 IAN BEVERIDGE
References
Bayuis, H. A. & Lane, C. (1920) A revision of the
Nematode family Gnathostomidae. Proc. Zool. Soc.
Lond. 245-310.
Beveripac, |, (9&5) A redeseription of Echinocephalus
uncinatus Molin, 1858 (Nematoda: Gnathostomatoidea)
from european rays, Dasyatis pastinaca (Linnaeus
1758). Aull. mus, nat. Hist, Paris, 4 me sér. 7, 762-780.
Drarporrr, T, L., Brooxs, D. R. & THorson, T, B,
(1981) A new species of Echinocephalus (Nematoda:
Ganthostomatidae) from neotropical stingrays with
comments on Ff. diazi. J. Parasitol. 67, 433-439.
& Ko, R. C. (1983) Echinocephalus overstreeti
sp.n. (Nematoda: Gnathostomatidae) in the stingray,
Taeniura melanospilos Bleeker, from the Marquesas
Islands, with comments on £, sinensis Ko, 1975. Prac.
Helminthol. Soc. Wash, 50, 285-293.
Hooper, J. N. A. (1983) Parasites of estuarine and
oceanic flathead fishes (family Platycephalidae) from
northern New South Wales. Aust. J) Zool. Suppl. Ser.
90, 1-69.
JOHNSTON, I. H. & Mawson, P. M. (1941) Nematodes
from Australian marine mammals, Rec. S. Aust. Mus.
6, 429-434,
& (1943) Some nematodes from Australian
élasmobranchs. Trans. R, Soc. S. Aust, 67, 187-190.
& (1945a) Parasitic nematodes, Rep. Brit.
Aust, New Zeal, Antarct. Res. Exped. 1929-1931, B, 5,
1-141.
&— — (1945b) Some parasitic nematodes from
South Australian marine fish. Trans. R. Soc. S. Aust.
69, 114-117,
Ko, R. C, (1975) Echinocephalus sinensis n. sp.
(Nematoda: Gnathostomatidae) from the ray
(Aetobatus flagellum) in Hong Kong, Southern China.
Cun. J. Zoal. 53, 490-500,
Lestcr, R. J. G. Biair, D. & HFAio, D. (1980)
Nematodes from scallops and turtles trom Shark Bay,
Western Australia. Ausr, J, Mar. Freshwater Res. 3A,
713-717.
McLauGuiin, R. H. & O’Gower, A. K. (197L) Life
history and underwater studies of a heterodont shark.
Ecol. Monog, 41, 271-289.
MiLLEMAN, R. E. (1951) Echinocephalus pseudo-
uacinatus n, sp, a nematode parasite of the abalone.
J, Parasitol, 37, 435-439.
(1963) Studies on the taxonomy and life history
of echinocephalid worms (Nematoda: Spiruroidea) with
a complete description of Echinocephalus pseudo-
Hucinatus Milleman, 1951_ Ibid 49, 754-764.
Soota, T. D. (1983) Studies on nematode parasites af
Indian vertebrates. I. Fishes. Rec. Zool. Survey India.
Mise. Publ. Ove. Pap. no. 54.
Troncy, P-M. (1969) Description de deux nouvelles
especes de nématodes parasites de poissons. Bull. mus.
nat. Hist. nat, Paris, 2 éme sér., 4, 589-605.
FOSSIL BROOD CELLS OF STENOTRITID BEES (HYMENOPTERA:
APOIDEA) FROMTHE PLEISTOCENE OF SOUTH AUSTRALIA
BY TERRY F. HOUSTON*
Summary
Calcareous fossils from the west coast of Eyre Peninsula, South Australia, previously recognized
correctly as petrified brood cells of burrowing bees and denoted by the ichnospecies names
Celliforma bedfordi and C. septata by Zeuner & Manning (1976), are assigned to the bee family
Stenotritidae. Notes are provided on a fossil site and samples from it are described, figured and
discussed.
KEY WORDS: fossils, brood cells, stenotritid bees, Pleistocene, South Australia.
FOSSIL BROOD CELLS OF STENOTRITID BEES (HYMENOPTERA: APOIDEA) FROM
THE PLEISTOCENE OF SOUTH AUSTRALIA
by TERRY F, HOUSTON*
Summary
Houston, T. F (1987) Fossil brood cells of stenotritid bees (Hymenoptera: Apoidea) from the Pleistocene
of South Australia. Thins. R. Soc. S. Aust, 111(2), 93-97, 29 May, 1987.
Calcareous fossils from the west coast of Eyre Peninsula, South Australia, previously recognized correcly
as petrified brood cells of burrowing bees and denoted by (he ichnospecies names Celliforma bedforai
and C. septata by Zeuner & Manning (1976), are assigned to the bee family Stenotritidae, Notes are provided
on it fossil site and samples from it are described, figured and discussed,
Kev Worbs: fossils, brood cells, slenotrilid bees, Pleistocene, South Australia.
Introduction
The fossils forming the subject of this paper were
first described by Zeuner & Manning (1976) who
recogmzed them as the petrified brood cells of
burrowing bees but could not identify them further.
Instead, they referred them to the ichnogenus
Celliferma recognizing two ichnospecies C bedfordi
and C, seplata, Zeuner and Manning’s specimens
were collected from “coastal (travertine and
consolidated dune-rock"” of Pleistocene or Sub-
Recent age at Venus Bay on the west coast of Eyre
Peninsula, South Australia, and are lodged in the
British Museum of Natural History, Their descrip-
tion of the fossils contains some confusing errors
and these are dealt with tater.
Additional specimens of the fossils from the west
coast of South Australia were located in the South
Australian Museum (registered numbers SAM
P24877-82). They were collected by A, Crooks tram
“S ml [miles] east of Rocky Point". Guided by
directions from this collector, 1 was able to locate
a bed of the fossils in coastal cliffs 0.5 km west of
Scott Point (32°01S, 132°23'E) and abour 6 kin
west of Fowlers Bay. | observed the form and
distribunon of the fossils and obtained samples for
the Western Australian Museum (registered nos.
WAM 86.723-86.730) on 7 January 1985,
Recent studies af stenotritid bee nests (Houston
1984; Houston & Thorp 1984) have revealed several
features which distinguish stenotritid broad cells
fram those of other bees, These same features
characterise the fossils known as Cel//forma septata
and ©. bedfordi and are the basis of my contention
that the fossus are a legacy of the nesting activities
of stenotritid bees,
* Western Australian Museum, Francis Street. Perth,
Western Australia 6000.
' Flint, R, B. (1986) Explanatory Notes, Nuyts (:250 000
Map Sheet area. South Australian Department of Mines
and Energy report 86/7 (unpublished).
Observations
Description of fossil site
The geological terminology employed here
follaws Flint (1986).!
The bed of fossils found near Scott Point is
located in an embayment of the coastal cliffs and
forms a band about 20 om high and approximately
30 m Jong in the cliff face (Fig. 1). The fossil band
rests ona hard calerete shelf that rises gently at each
side and beneath this are three other calcrete shelves
separated by sott aeolian calcarenite. The fossil
band appears to be situated in the lowest horizon
of a fossil colluvial soil about | m high, Three
horizons are evident: (J) an upper zone of brown
silt about 30-40 em high (presumably rhe original
surface); (2) a middle zone of calerete breccia; and
(3) a lower zone of pale yellow clayey calcareous
aeolianite in which the fossils are embedded. Above
this fossil soil is deep white aeolianite capped by
calerete and on this are recent acolian dunes
(suffering deflatian),
The fossil brood cells are very numerous and
aceur densely packed, some being welded together
or to the calcrete basement. All are horizontal or
subhorizontal.
Searches nearby located only a small number of
Other brood cells welded to a lower calerete shelf,
Description af fossils
Loose fassils fram the Seatt Point site (Fig. 2)
measure 40-70 mm in fength and 17-20 mm in
diameter. Usually they are slightly curved, rounded
at one end and truncated or concave at the other,
Within the rounded end of each is a somewhat
ovoidal chamber (either empty or soil-filled)
measuring 13 mm in maximum diameter and, when
plugged, 28 mm in maximum length. When nar
plugged, the chamber is continuous with 4
cylindrical burrow which extends ta the truncate
94 T. F. HOUSTON
Fig. 1. Fossil site west of Scott Point (background). Fossiliferous horizon in upper Bridgewater Formation is arrowed.
Fig. 2. Loose fossils from upper Bridgewater Formation west of Scott Point. Holes in the specimens on the right
were probably made by emergence of a parasite (upper) and a bee occupant (lower). (WAM 86.723-86.726).
FOSSIL BROOD CELLS OF BEES 95
end. The cemented wall of the chamber varies from
2-7 mm in thickness and while variably rough
externally it is quite smooth internally (Figs 2, 3).
Many of the fossils are sealed with a complex
closure, In the neck of each chamber is a plug about
10 mm long, the inner face of which usually exhibits
an inverted conical spiral pattern and the outer face
being smoothly concave. Between this and the
truncate end may be 4-8 thin concave cemented
partitions, 3-4 mm apart. The spaces between them
are solidly filled with fine soil and gravel particles
up to 5 mm across.
Figs 3 and 4. Sagittal section of an intact fossil (WAM
86.727) from west of Scott Point (3) and interpretation
of its structure (4). Legend: ch, cell chamber; cs, concave
septa (Ist-3rd appear to have been disturbed in lower
parts); g, gravel particles; p, cell plug.
Other fossils lack the plugs and partitions and
are uniformly filled with soil and gravel. Many
specimens with intact closures had a lateral hole
9-10 mm in diameter about 30 mm from the
rounded end while a few had smaller holes in a
similar position (Fig. 2).
Amongst the S. Aust. Museum specimens are
some that are embedded in a solid calcrete rock
matrix (e.g. P24879 and P24882) and which are
evidently older than those described above. Never-
theless, they clearly show the series of concave
partitions in the chamber access burrows and their
internal dimensions match those of free fossils
(Fig. 5).
Discussion
An uncritical observer could easily confuse the
fossils described here with other more common
Fig. 5. Trace fossils embedded in solid calcrete from lower
Bridgewater Formation, 8 km west of Rocky Point.
(SAM P24879, P24882). Scale lines, 1 cm.
kinds of fossils derived from the pupal cases of
Coleoptera, in particular the large weevils,
Leptopius spp. These were described and figured
by Lea (1925) and are distinguishable by their more
ovoidal form (both externally and internally), the
absence of a chamber closure, and usually by their
larger diameter.
That the fossils described here are the petrified
brood cells of a stenotritid bee can scarcely be
96 T. 0, HOUSTON
doubled in view of their close resemblances to thre
brood cells of extant species. Diagnostic features
of srenarritid cells (Houston & Thorp 1984) which
are observable in the fossils are: (1) built-in walls
of cell chamber continuous with those of access
burrow: (2) cell plug with concave spiral pattern on
inner face, outer face smooth and concave: (3)
aceess burrow sealed with one or more convave
cemented partitions; (4) gravel particles included in
soil-filling between plug and partitions; and (5) cells
more ov jess horizontally orented,
Only two genera of Stenatritidae, Stenairitus and
Ctenocolletes, are recognized and our present
knowledge af stenotntid nest architecture provides
no means of distinguishing the braod cells af these
two taxa. However, adults of Ctevrocol/etes are
larger on average than those of Stenosritus and,
since the internal dimensions of the fossil cells
marginally exceed those of the largest known
Clenocolletes cells (C michalsoni (Cockerell);
unpublished personal observations), it seems more
probable that they are derived from the construc-
tions Ol a Crenocalletes species. Were that species
to be extant, it would most probably be C
Julveseens Houston, the only stenotriod know to
inhabit. the country Fringing the Great Australian
Right, Unfortunately, neste of this species are
unknown, One of the extraordinary features of the
fossils is the high pAumber of concave septa
occurring in the cell clasures (up to eight of them),
Three is the maximum number chserved in the
closures of extant Stenotritidae (Houston & Thorp
1984),
Although Zeuner & Manning (1976) recognize
two ichmospecies of fossil cells from Venus Bay,
ihere is n0 reason to suppose thar mare than one
hee species was Invalved in the making. Celliforena
bedfordi was distinguished by the cell plug having
a concave spiral pattern on its inner face (absent
im C. septafa), The spiral pattern would be typical
al cells which remained undisturbed after closure
(presumably when occupants bad died). The-closure
of cells from which adults had emerged would be
modified or destroyed as the becupants burrowed
oul.
Retallack (1984) hkened C bedfordi and ©
septate to the earthen braod cells of living Melitorma
and Plilethrix bees (Anthophoridae) but the
resemblatices are few and are far surpassed by those
of stenotritid brood cells.
The stenotritid fossils occur Ig packs forming part
ofthe Bridgewater Farmatian which was laid down
during the middle Pleistocene (Flint 1986),! Fossils
occurring inthe lower part of this formation (those
embedded in solid calerete) are of uncertain age but
possibly as old as 700 (00 years while those from
(he upper part (the separable or only slightly fused
fossils) may be up to 100 000 years old (R. B. Flint,
personal communication).
J consider the fossil bed in the upper Bridgewater
Formation near Scott Point to represent a perennial
nesting site where hundreds of female bees over
numerous generations had nested gregariausly (bul
individually) in colluyial soil filling a limestone
swale. Stenotritids burrow into level ground wid
construct broad cells at the lower ends of the shafts,
They cement the earthen walls of the cells and the
access burrows wilh some secretion so thal they are
quile durable constructions which may remain for
years. There must be a tendency for lime to be
deposited in the walls of the cells ar a greater rate
than in the surrounding soil, Thus they become
calcified first as separate entities, then gradually
fused to one another, then finally encased in a solid
calerele matrix,
Because of the high frequency of unperforated
closures amongst the specimens examined it must
be supposed that either there was a high mortality
rate amongst immatures or that emerging adults by-
passed the closures. Evidence for the latter
possibility occurred in the many specimens with a
large lateral hole in near proximity to the cell plug
(Fig, 2). However, such lateral emergence is so far
Unknown amongst extant Stenotritidae and is
atypical of bees generally, emergence usually
occurring via the old access burrows and involving
demolition of the closures,
Comment
The following corrections are made to Zeuner & Manning
(1976).
Page 205: Under "Remarks" for €. bedfordi, the second
sentence should be resiructured to make it clear that the
Whole fossils are “some 6-7 cm long by 2. cm in diameter”,
nol the scptate cross walls.
Registration number for paralype of C. sepra/a should be
tn. 31433 (not 34133).
Camion for Plate 1, Fig, 9, should read Paratype (mor
Holotype).
Plate | figure numbers should he altered as follows: 6 to
9.7 to 6, and 9 tu 7
Caption for Plate 2, Pig. 1, should read Holotype (noc
Paratype).
Acknowledgntetits
Preparation of this paper was facilitated by the
generous assistance of a purnber of people. Ir
particular [ wish to thank the following: Messrs
Alistair Crooks and Richard Flint (Department ef
Mines and Energy, South Australia) and the
Director-General of their department for informa-
tion on fossil sites in South Australia and access
FOSSII. BROOD CELLS OF BEES 97
to an unpublished report; Mr Tony Gayski (Geology
Department, University of Western Australia) for
preparing sections of the fossils; Dr Ken McNamara
(Department of Palaeontology, Western Australian
Museum) for providing copies of some relevant
literature; and Mr Neville Pledge (Department of
Palaeontology, South Australian Museum) for
providing access to specimens and data in his
department. Dr McNamara and Mr Pledge also
read a draft of this paper and made useful
suggestions for its improvement, as did two referees,
Dr B. Webby and Professor A. Seilacher.
References
Houston, T. F. (1984) Biological observations of bees in
the genus Crenocolletes (Hymenoptera: Stenotritidae).
Rec. West. Aust. Mus. 11(2), 153-172.
& THorP, R. W. (1984) Bionomics of the bee
Stenotritus greavesi and ethological characteristics of
Stenotritidae (Hymenoptera). [bid. 11(4), 375-385.
Lea, A. M. (1925) Notes on some calcareous insect
puparia. Rec. S. Aust. Mus. 3, 35-36.
RETALLACK, G. J, (1984) Trace fossils of burrawing
beetles and bees in an oligocene paleosol, Badlands
National Park, South Dakota. J Puleont. §8(2), 571-592.
ZEUNER, F. E. & MANNING, F. J. (1976) A monograph on
fossil bees (Hymenoptera: Apoidea). Bull, Br. Mus. nat.
Hist, Geology 27(3), 149-268.
ANOPLOZETES, A NEW GENUS OF ZETOMOTRICHIDAE
(ACARIDA:CRYPTOSTIGMATA) FROM SOUTH AUSTRALIA
BY DAVID C. LEE & GEORGE A. PAJAK*
Summary
Anoplozetes jamiesoni gen. nov., sp. nov. is described from arid tussock grassland in the Victoria
Desert, northern South Australia. The Zetomotrichinae are considered and a key provided to
separate the seven genera. This is the first record of Zetomotrichidae from Australasia.
KEY WORDS: Acarida, Zetomotrichinae, new family record, Anoplozetes jamiesoni, new genus,
new species, South Australia.
ANOPLOZETES, A NEW GENUS OF ZETOMOTRICHIDAE (ACARIDA:;
CRYPTOSTIGMATA) FROM SOUTH AUSTRALIA
by Davin C. LEB & GEORGE A. PAJAK*
Summary
lan, DC. & Paiak, G. A. (1987) Anoplozeres, a new genus of Zelomotrichidae (Acarida: Crepiostigmata}
from South Australia. Trans. R. Soc, S, Aust, LL(2), 99-403, 29 May, 1987.
Anouplogeles jamiesoni gen. nov.,.sp. nov. is described from arid tussack grassland in Lhe Victoria Desert,
northern South Australia. "he Aetamotrichinac arc considered and a key provided to separate the seven
genera, This is the first record of Zetomotrichidae from Australasia,
Key Worbs; Acarida, Zeromotrichinag, new family record, Anop/ozeles samiesoni, Tew Bens, new
species, South Australia.
Introduction
This publication is part of an ongoing study (Lee
HORI; 1982; (985; in press) of sarcoptiform mites of
South Australian soils, sampled from nine Mlorally
diverse sites. The new species described here was
collected only at the arid grassland site. It is
established as the type of a new genus and requires
modification of the subfamily diagnosis. The
Zetomotrichidae include two subfamilies of which
the Rohriinae Balogh & Balogh, 1984 from Brazil
are not considered. The briefness of the description
of the single species of Rohriinac makes it uncertain
as lo whether or mot some of the diagnostic
character states of the Zetomotrichinae should
apply to the whole family.
Materials and Methods
The notation and methods of measurements
follow Lee (1981) with modifications made by Lee
(in press), Measurements are in microns (jm). The
crochantera are illustrated (Fig. 3), although normal,
lo emphasize their similarity to both those of the
short-legged Constrictohares (Lee in press) and the
saltatory Zefomoirichus, The mites examined are
deposited in the South Australian Museum.
ZETOMOTRICHINAE Grandjean
Yetomotrichidae Grandjean, 1954: 16,
Diognosis: Comalida, Planotissurae, Poronotae.
Onpodoidea. Zetomotrichidae. Noral foramina
absent, but numerous scattered refractile micro-
pores, ‘Soma spindle-shaped, dorsosejugal furrow
“ Div. of Natural Sctence, South Australian Muscum,
North Terrace, Adelaide, S, Aust, 5000.
mainly absent, row of sigilla across line it would
occupy. Rostral margin of prateronotum
denticulate. Lamella (seta 21 to 72) absent, Ptero-
morphs absent but conspicuous tooth-shaped
provess bearing seta (ZL) on hysteronotal shoulder
(both directed forward) and deep limbus around
lateral and posterior hysteronotal margin. Hystero-
netum with 10 pairs (3.7, 52, 2.9) of setae. External
malae narrow, not ventrally obscuring oral setae,
Legs long, leg [V longest (femur-tarsus longer than
half somal length), both tibia and tarsus 1V
subequal in length to femur 1V, femur J and LI with
long stalk, tarsi I, 11, WE narrow proximally
(subequal to distal diameter), pretarst pedunculate
with three claws,
Distribution: Previously known from two main
areas: around the Mediterranean, Caspian scas and
in India, and from the Andes in Peru and southi-
ward, Particular species recorded from caves,
tussock grass or as saxicolous, but also known from
woodland and forest litter,
Remarks: The Zetomotrichidae are unique in the
Oripodoidea (-Oribatuloidea; Balogh & Balogh,
1984, sce Lee in press), and unusual in the Poro-
notae, in Jacking foramina. Although the refractile
notal micropores may serve the same function, they
do not appear to be homologous. Despite the
absence of this diagnostic character State, the only
nymphs described (Covarrubias 1969) belong to the
"Excentrosclerosae” (=Oripodoidea, see Lee in
press).
Balogh & Balogh (1984) record only three genera
in the Zetomotrichinae and incorrectly date the
authority of the hame-as “1934”. Besides the new
genus established in this paper, there are six genera
recorded here in chronological order of establish-
ment date, with references to fuller deseriptions for
two genera, as well as number of species and
distribution:
1H! D.C, LEE & G. A. PAJAK
Zelvorwirichus Grandjean. 1934: One species;
Algeria (Pm), ? Caucasus (Vs), Pakistan (Oi)
Mikizeres Hanwner, 1958 (Covarrubias, (969
includes onty description of immatures for the
family): Two species: Aigentina, Peru and Chile
(NTc).
Ghifarevus Krivoluisky, 1966 (Subias & Pérez-Iniga,
1977): Three species; Spain (Pm), Uzbekistan and
lurknienia in Central Asia (Ps near Pm),
Paliliducarus Krivolutsky, 1974: Qne species;
Turkmenia in Central Asia (Ps near Pm),
Oelasacarus Bernini, 1978; One species;
Monitcerista Island (Pm),
Kevuloiricdus Mahunka, 1985: One spectes; South
India (Oc).
KEY TO ZRTOMOTRICHINAE GENERA
(Adults)
b, Noral setae Z1 & 72 similar to 42_ Pores Aj2, fy'S slit-
like, enlarged, twice length of setit 22. No pyriform
organ itt drea bf pore Af3, no himetal organ or
subapical pracess present, Coxisternal seta (2 & 72
more than twice length of MW & W. Genital shields
bearing fuur setal pairs neh two pairs of parainal
setae (25a) , ' Anoplaretes
Notal setuc 2 d& j2 at jest twice ap long and stout
a3 72, 71 usually not setose, Pores A/?, A/3, uf similarly
slit-like, subequal in length (0 or shorter thar seta 22.
Humeral organ or subapical process present, pyrite
organ may be present in area of pore 4/2. Coxisteraal
sctae /2 & #172 subequal in lengih to or sharter ahan
AL & ML. Genital shields with three ta five setal pairs,
cwo or three pairs of paranal setae, but never Vee,
28d
2. Hysteronot#l pores 4/3, A/6 slit-like, subequal mm lengi
(4 seta Z2. Covisiernal setae /2 & 12 suboqual in
tenerh tn J) & 7/1. Rostral margin denticulure, teerh
shape similar, Jarger lowards centre ,,,, 0 ...... 3
Ilysteranonal pore 4/6 slit-like, 4/3 invouspiewous or
absent, pyriform onan in similar location. Coxisternal
setac 72 & (7/2 less than half tength of entarged f1 &
{7}, Rostral murgin with convey apes borderod by large
feetly, , .- 100 bapececcere tihheess sven vere &
3. Genital and paral setal formula 4/42, Wee. +
Genital and paranal seral foruiula $JZy, 28a... 5
4. Hiimeral process present, oo humeral organ. Soma
yellow brown, integumear withowr linear sculpturing.
Kosteal woth larger towards centre, .... Ghilarovus
Humerul process absent (humeral organ not known).
Soma pale yellow, Integament- covered in parallel linear
sculprurine, Kostral teeth subequal in size Pallidacarus
3. Hinicral organ without associated larger sacculate
SEPLICIUTE eve APs . Atikizetes
Humeral argan with associated larger sacculire
HEMOTUINE ye chen Oxglasaearis
6, Genithl setal formiute 3/%2, Hysteronotal seta Zt
subeqital In length to el. Liner hysteronatal, region
framed by conspicuous line (adaxial to setae, 71, 22;
24, 35, 25, 16) , Kerulotrichus
Cienital seta) formula 4/Ze, Hysteronotal seta /4
shiurter than st, No conspicuous line fraraing part of
hysteronotum, . . Ly. Zelamotricins
The short descriptions of Pallidacarus and
Keralotrighus were a drawback in constructing the
key. Pallidacarus 1s assumed lu be similar to
Ghilerovus, but it would be useful to know whether
or not it has a humeral organ. Aeralorrichus can
be delineated from Zefurmotrichus, but there is a
suggestion in an illustration (Mahunka 1985;
Fig, 42) that it may have acetabulum IV similarly
dorsal to acctabulum [11, and it is, therefore,
assumed here that it has character states as for
Zetamatrichus which are associated with jumping,
such as The enlarged dorsal selge on tarsus IV, The
homologies of the pyriform organ, humeral organ
and pssociated sclerire or sacculate structure nexl
ta be clearly established, The phylogenetic model
held in this study is that Anop/ozetes is prinmtive
and Aerulorichus and Zefumotrichus are the latest
derived sister group, The remaining gener
apparently form ad intermediate group, amonast
Which the similarity heiween the South American
Mikizetes and the others, which are all Palacarctie
genera, suggests that this group is, or has been,
widespread. The loss.of setuc is not valuable as an
indicator of derivation, different losses on the venjer
not being correlated, and setal losses are only
ovcasional in the leg chaetoraxy (Ghilarovus, only
four setae on femur IW; Zeremeotrichus, only two
setac on tibia I).
re
Anoplozetes gen, nov,
Type species! Anoplozetes jamiesoni sp. nov.
Diagnosis: Zctamotrichinae: Notal setae Zi) and 2
fing, setose, similar to 72, Hysteronotal pores Af
and Asa slit-fike, enlarged, twice length of seta 72,
Pyriform ora in area of pore 4/3 wbsent. Humeral
organ and subapical humeral process absent. Coxi
sternal setae 72 and 7772 at least twice length of SI
and ///1. Genital shiclds bearing four setal pairs
(4/22). Two pairs (2S) of paranal setae. Tarsus 1Y
without enlarged dorsal setae.
Remarks: Two. character stales previously diagnostle
of Zelomoatrichinae are not represented, These are
the enlarged setac Zl and /2 and the presence of
cither hurneral organ or subapical process. Also, the
derived stutes of the type-genus (and. possibly
Keralotrichus) associated wilh jumping, such as the
presence of enlurged dorsal setag on tarsus TV and
the positioning of acetabulum LV dorsal to aceia-
bulom III, are absence. On the other hand, although
ANOPLOZETES, ZETOMOTRICHID MITE 101
the lack of these states has been regarded (Covar-
Tubias 1969; Bernini 1978) as indicating a lack of
adaptation of Jeg |V for jumping, it is possible that
the unusual length of leg 1V may be related to some
ability to jump. Anoplozetes, lacking the derived
character states of the hysteronotal shoulder and
leg IV, is considered to be the most primitive genus
in Zetomotrichinae,
Anoplozetes jamiesoni sp. nov.
FIGS 1-3
Female: General appearance shiny yellow-brown,
cerotegument inconspicuous, most somal setae fine
and short, Integument generally smooth, irregularly
placed refractile micropores on notum, few fine
striations around proteronotal seta j2 and
reticulations on coxisternum. Idiosomal length 323
(4, 298-344). Appendage lengths (femur-tarsus, for
333) — I 173, 11 150, 111 149, 1V 202; tibial height
— 1 23, II 16, 111 13, [V 16,
Prosternum with deep mentotectum, crossed
laterally by longitudinal ridge. Custodium extending
forward from pedotectum II, no discidium nor
circumpedal ridge present. Five refractile ridges
running vertically up into pleural region from
between aceltabulae [I1/1V. Coxisternal sctae in 10
pairs (3/1/, W//, 34/1, 31), lateral setae longer, seta
100gLm
Figs. I-2: Aneplozetes jamiesoni sp, nov. Fig. 1, notum of soma. Fig. 2, sternum of soma.
102 D.C. LEE & G, A, PAJAK
M2 and [772 more than twice length of Ji and
iN.
Proteronatal rostral margin weakly denticulate,
laterally extending backward behind level of scta
JJ (not illustrated because pointing ventrally).
Without lamella between setae zl and 22, although
short apodeme anterior to 72 and weak ridge behind
zl. Proteronotal setae in five pairs: /l, zl long and
stout, /l with conspicuous unilateral row of cilia,
z2 long with conspicuous bilateral rows of cilia, j2
and s2 fine and short, with s2 positioned ventrad
to. bothridium (around 22),
Opisthosternal shield margin (Fig. 2, broken line)
extending unusually far behind anal shields and
marginally overlapped by hysteronotal limbus.
Chaetotaxy: 4/Zg, |Se, 2JZa, 2Sa. On genital
shield, anterior setae Jonger (han posterior setae.
Slit-like pore Saf almost transverse.
Hysteronotal margin with forwardly directed
triangulate shoulders, elsewhere ventrally directed
limbus overlapping margin of opisthosternal shield
and with two posterior hyaline lobes, one over-
lapping the other (note Fig. 2 has no representation
of hysteronotal margin with unsclerotized cuticle
between it and opisthosternal shield). Chaetotaxy:
34, 5Z, 2S. Two pairs of slit-like pores, Af3 and A/6,
over [hree Limes length of nearby setae, 4/4 and hf5
subequal in length to such setae (not completely
visible from above, see Fig. 1). Many refractile
micropores scattered over surface.
Legs long, order of decreasing length IV, I, IL,
Il, leg LV (femur-tarsus) longer than half somal
length, Femora I and I! with long stalk, short
ventral flange bearing ventral seta on femur II.
Femora IIf and 1V with short stalk, caput large.
rectangular, anteroposteriorly flattened with shallow
100m
a
m mw
hig. 3. Anaplozetes jamiesoni yp. nov., posterior aspect of right legs, showing only flagelliform solenidia and one
ventral seta.
ANOPLOZETES, ZETOMOTRICIUD MITE 103
ventral flange. Tarsi long (J-1.5» length of tibia),
anterior three tarsi gradually tapering proximally
to Jess than quarter of breadth. One long Slagelli-
form solenidium on tibia [, If and tarsus 1, other
solenidia setiform or bacillitorm and shorter (not
illustrated, Fig, 3). Solenidiotaxy: 1(1,2,2,), 11(1,1,2),
Hi001,1,0), £¥(0,1,0),
Length of finely wrinkled ovipositor tube, 129
(soma 333), including three lobes (23).-Ovipositor
bearing 16 setac, subequal in size tu each other, all
longer than coxisternal seta #2, proximal setae (pg)
with lips reaching bases of distal setae (dg). Only
one female with a single large oval eyg (141 « 102),
surface smuoth. Boli may be granular, including
amorphous. fragments, non-septate hyaline tube
fragments (?hyphae), regular minute rods (? bacilli)
or spheres (?spores), and sometimes larger fungal
(Deuteremycotina, 2Alternaria) conidia (one bolus
with 26 dark brown septate conidia, 18-26 long).
Male: As female except for measurements and
spermapositor, ldiosomal length 289 (14, 258-314).
Spermapositor very short, length less than greatest
breadth (24), anierior-posterior axis of included
bilobed sclerite 22. Spermapositor bearing 14 setae,
shart, subequal to each other, length about twice
diameter of setal base:
Material examined: Holotype female (N1986244),
three paratype females (N1986245-N1986247), 14
paratype males (N1986248-N1986261), bases of love
grass (fragrostis eriopoda) tussocks, near Emu
{28°41'S, 132°O8’E), |(1.x,1974, D. C. Lee,
Distribution: Australia (Aa). South Australia, Great
Victoria Desert, tussock grassland, 4 females, 14
males, in 5 of 8x25 cm* samples.
Remarks: This. species, being the type of a
monotypic genus, has diagnostic character states as
tor Anuplozetes, There is. an unusually high propor-
tion of males possibly refleciing an adaptation to
the environment or that the material collected does
Nol represent a vormal population,
The species is named after Dr B. J. M. Jamieson,
Queensland University, for his encouragement
during the earlier slages of this project on soil mites.
Acknowledgments
We wish to thank the Australian Biological
Resouces Study for a grant to one of us. (D,L,)
funding the salary of the other (G.P.) and Ms Jenni
Thurmerc for vie woration and presentation of the
hyures.
References
BavocH, J, & BAtocH, P, (1984) Review of the
Oribatuloidea Thor, 1929 (Acari; Oribatei). dere goat,
Aung, 30, 257-313.
BERNINI, FP. (1978) Notulae Oribatologicae XVIIL.
Oglasacarus oglasae i. gen., 1. sp., Un nuovo
Zetomotrichidae raccolto sull'isola di Montecristo
(Acarida, Ovibatida), Redia 61, 273-289,
COVARRUBIAS, R, (1969) Observations sur le gerire
Mikizeley (Oribates). Acarologia 11, 828-846,
GRANDIFAN, F. (1934) Oribates de l'Afrique du Nord (Ze
serie), Bull, Sac, Ais Nat, Afr. Nord 25, 235-252.
(1954) Zeromuitrichns lacrimuns, Acarien sauteur
(Oribate) (Avar, Zetomotrichidae) Ann. Soc. Entomol.
France 123, 1-16.
HAMMER, M, (1958) Investigations on the Oribatid faund
of the Andes Mountains. I, The Argentine and Bolivia.
Biol. Skr. Dan. Vid. Selsk. 10, 1-129.
Krivyouersky, D, A. (1966) Q pantsimyh klechtchah
{Oribatei, Avariformes) poich srednei Asii. Zool Zn
45, 1628-1638,
(1975) Zetomotrichidac, Jn Ghilarov M. S. &
Krivolutsky D. A. (Eds), Opredelitel obitaiouchi& ¥
potchve Klechtchi. (Nauka, Moscow).
Lee, D, C. (1981) Sarcopliformes (Acari) of Soattr
Australian soils, 1. Notation. 2. Bifemoraia ane
Ptyctima (Cryptostigmata). Rec, 8. Aust Aqus UR,
199-222.
(1982) Sarcoptiformes (Acari) of South Australian
soils. 3. Arthronotina (Cryptostigmata). Ibid 1s,
327-359,
(1985) Sareoptiformes (Acari) of South Australian
soils. 4. Primitive oribate mites (Crypitoatigmata) with
an extensive, unfissured bysteronoral shieltl ana
aptychoid. bid, 19, 39-67,
(in press) Introductory study of advanved oribate
mites (Acaridia: Cryptostigmata: Planofissurae) and a
redescription of the only valul species of
Consrrictobates (Onipodoides). fbid, 21.
MAHUNKA, 8. (1985) Neve und interessante Milben aus
dem Genfer Museum. .1V. Oribatids from South india
| (Acari: Oribatida), Revue suisse Zuv7 92, 367-383,
Supias, L. S. & Peere-Iriae, €. (1977) Notes sur les
Oributes U'Espangne. 1. Description de Chilarowes
hispanicus n. sp. et quelques considérations suc tes
eetomatsgetane (Acari, Oribatei), Acarologia 1&8,
729-739,
THE SUBORDER CHORIPLACINA STAROBOGATOV & SIRENKO, 1975
WITH A REDESCRIPTION OF CHORZPLAX GRAYZ (H. ADAMS &
ANGAS, 1864) (MOLLUSRA:POLYPLACOPHORA)
BY K. L. GOWLETT-HOLMES*
Summary
The suborder Choriplacina Starobogatov & Sirenko, 1975 is reevaluated and redefined as a
suborder of the order Neoloricata. The sole living member of this suborder, Choriplax grayi, is
redescribed and its habitat discussed. The family Afossochitonidae is reyiewed and placed in
synonymy with the Acanthochitonidae. Afossochiton is placed in the Acanthochitonidae. Lirachiton
is placed in synonymy with the subgenus Bassethullia of Notoplax in the Acanthochitonidae.
Glyptochitonidae is recognised and contains the genus Glyptochiton.
KEY WORDS: Chiton, Polyplacophora, Choriplacina, Choriplacidae, Choriplax grayi,
Afossochitonidae, Acanthochitonidae, Glyptochitonidae.
THE SUBORDER CHORIPLACINA STAROBOGATOV & SIRENKO, 1975 WITH A
REDESCRIPTION OF CHORIPLAX GRAYI (H. ADAMS & ANGAS, 1864) (MOLLUSCA:
POLYPLACOPHORA)
by K. L. GOWLETT-HOLMES*
Summary
Gow.ett-Hoimes, K. L. (1987) The suborder Choriplacina Starobogatov & Sirenko, 1975 with a
redescription of Choriplax grayi (H. Adams & Angas, 1864). (Mollusca: Polyplacophora). Trans. R. Soc.
S. Aust, 111(2), 105-110, 29 May, 1987,
The suborder Choriplacina Starobogatov & Sirenko, 1975 is reevaluated and redefined as a suborder
of the order Neoloricata. The sole living member of this suborder, Choriplax grayi, is redescribed and
its habitat discussed. The family Afossochitonidae is reviewed and placed in synonymy with the
Acanthochitonidae. Afossochiton is placed in the Acanthochitonidae. Lirachiton is placed in synonymy
with the subgenus Bassethullia of Notoplax in the Acanthochitonidae. Glyptochitonidae is recognised and
contains the genus G/yptochiton.
Key Worps: Chiton, Polyplacophora, Choriplacina, Choriplacidae, Choriplax grayi,
Afossochitonidae, Acanthochitonidae, Glyptochitonidae.
Introduction
The suborder Choriplacina was erected by
Starobogatov & Sirenko (1975) to accommodate the
two monogeneric families, Glyptochitonidae
(Carboniferous) and Choriplacidae (Recent).
However, both families were regarded as synonyms
of Afossochitonidae by Van Belle (1975, 1981, 1983),
Ferreira (1981) and Kaas & Van Belle (1985), who
placed them in the suborder Lepidopleurina, based
on characters of the shell (the animal of the
Choriplacidae being then unknown).
Starobogatov & Sirenko (1975) placed Choriplax
grayi in the Choriplacina because of its reduced
tegmentum, but Van Belle (1975, 1983) and Ferreira
(1981) placed this species in the Afossochitonidae
(Lepidopleurina) because of its unslit insertion
plates, regarding the reduced tegmentum as of
generic value only.
The recent discovery of several complete
specimens of C. grayi has provided the opportunity
to fully redescribe this species and reexamine its
systematic position. Consquently it is necessary to
reevaluate the suborder Choriplacina Starobogatov
& Sirenko, 1975, based on the characters of
Choriplax grayi (H. Adams & Angas, 1864), the
only known extant species.
The following abbreviations are used: AM,
Australian Museum, Sydney; BMNH, British
Museum (Natural History), London; NMYV,
Museum of Victoria, Melbourne; SAM, South
Australian Museum, Adelaide. Colour descriptions
follow Kornerup & Wanscher (1978).
* South Australian Museum, North Terrace, Adelaide, S.
Aust. 5000
Systematics
Class Polyplacophora Blainville, 1816
Order Neoloricata Bergenhayn, 1955
Suborder Choriplacina Starobogatov & Sirenko,
1975
Diagnosis: Small to large chitons with large unslit
insertion plates in all valves; tegmentum much
reduced; gills holobranchial.
Composition: This suborder contains the two
monogeneric families Choriplacidae and
Glyptochitonidae.
Family Choriplacidae Ashby, 1928
Diagnosis: Small to medium chitons with the
characteristics of the suborder.
Composition: This family contains only the
monotypic genus Choriplax, endemic to southern
Australia.
Genus Choriplax Pilsbry, 1894
Microplax H. Adams & Angas, 1864, p. 194 (Microplax
grayi H. Adams & Angas, 1864, original designation) not
Microplax Fieber, 1861 (Hemiptera).
Choriplax Pilsbry, 1894, p. 139 (Microplax grayi H.
Adams & Angas, 1864, original designation).
Diagnosis: Small to medium chitons; tegmentum
sculpture granular; articulamentum very large,
sutural laminae well developed, sinus obsolete;
girdle large and fleshy; gills holobranchial, abanal.
Composition: This genus contains the single species
Choriplax grayi.
Choriplax grayi (H. Adams & Angas, 1864)
FIGS 1-2
Microplax grayi H. Adams & Angas, 1864, p. 194; Angas,
1865, p. 58, pl. 2, fig. 16; Angas, 1867, p. 224; Pilsbry,
1892, p. 21, pl. 6, figs. 9-11.
106 K. L. GOWLETT-HOLMES
Fig. 1 Choriplax grayi. A. dry adult specimen, x3.2 (SAM D17443); B. dry juvenile specimen, x7 (SAM D16543);
C, median valves, x15 (BMNH 1877.11.7.2); D. posterior valve, x20 (BMNH 1877,11.7,2).
Choriplax grayi (H, Adams & Angas, 1864); Pilsbry, 1894,
p. 139; Ashby, 1924b, p. 383; Kaas & Van Belle, 1980, p.
56; Kaas & Van Belle, 1985, p. 204, fig. 95, map 16; Zeidler
& Gowlett, 1986, p. 105.
Lepidopleurus (Choriplax) grayi (H. Adams & Angas,
1864); Ashby, 1918, p. 83.
Choriplax grayi pattisoni Ashby, 1921, p. 137, pl. 9, figs,
la-c; Iredale & Hull, 1925, p. 100, pl. 11, fig. 28; Iredale
& Hull, 1927, p. 90, pl. 10, fig. 28; Kaas & Van Belle, 1980,
p. 56, 96; Zeidler & Gowlett, 1986, p. 105,
Choriplax grayi grayi (H. Adams & Angas, 1864); Iredale
& Hull, 1925, p. 99, pl. 1, figs. 24-27; Iredale & Hull,
1927, p. 89, pl. 10, figs, 24-27.
Choriplax pattisoni Ashby, 1921; Ashby, 1924a, p. 331;
Ashby, 1924b, p. 383; Cotton & Weeding, 1939; p, 188;
Cotton & Godfrey, 1940; p. 540; Cotton, 1964, p. 85, fig.
542, fig. 96.
Material examined
Types: Holotype (BMNH 1877.11.7.2) dredged from
Watsons Bay, Port Jackson, New South Wales,
collected by G. F. Angas (anterior valye missing,
CHORIPLACINA (POLYPLACOCOPHORA) 107
presumed lost). Choriplax grayi pattisoni: Holotype
(SAM _ D15019) from Cape Banks, South Australia,
collected by G. Pattison, 1918.
Other material: Tasmania: SAM D16542 (1) Fluted
Cape, Bruny Island, S. A. Shepherd, 11.ii.1972.
South Australia: SAM D17443 (1) Racecourse Bay,
Port MacDonnell, T. & M. Young, 1968;
SAM D16543 (2) Cape Northumberland, S. A,
Shepherd, 19.vii.1974. Western Australia:
NMV F51767 (2) Carnac Island, Perth, N.
Coleman, 1971; AM C151131 (1) Carnac Island,
Fremantle, N. Coleman, 18.xii.1971.
Species description: Small to medium chiton to 30
mim. Semicarinated; tegmentum discrete, very small;
more or less posteriorly positioned on valves.
Tegmentum colour variable: greyish-red to greyish-
orange to light brown. Articulamentum white with
pastel red under tegmentum and along posterior
edge of valves. Girdle very large, fleshy (Fig. 2A);
in dried specimens shrinking to thin horny covering
(Figs 1A, 1B). Girdle colour in living animal
matches tegmentum (Shepherd pers. comm.), dried
or alcohol preserved specimens greyish-yellow to
khaki.
Anterior valve tegmentum almost circular, with
slight beak; sculptured with longitudinal, granular
ridges posterior to beak, central area with radiating
granular riblets over 1/3 of area, granules becoming
coarser, then irregularly pustulose over remaining
area. Median valves (Fig. 1C) tegmentum heart
shaped, with distinct beak; sculptured like anterior
valve. Posterior valve (Fig. 1D) tegmentum tear-drop
shaped; antemucronal area with longitudinal,
granular ridges; mucro granulose, in anterior 1/3
of valve; postmucronal area like anterior valve
anterior to beak,
Girdle covers valves completely with transparent
layer over tegmentum; subsurface with granular
appearance; with sparse, 20-30 »m long, smooth,
conical, calcareous spicules, wider at the base in a
chitinous cup (according to Kaas & Van Belle 1985).
Gills (Fig. 2B) numerous, holobranchial, abanal,
23 to 28 medium to large ctenidia on each side.
Radula (Figs. 2C, 2D) with small weak central
teeth, heads weak and asymmetrical; first lateral
teeth broadly rounded, slightly narrower basally;
major lateral teeth with wide tricuspidate heads,
cusps sharp, of equal length.
Habitat: On the prostrate red alga Sonderopelta
coriacea Womersley & Sinkora, 1981; occasionally
on sponge on brown algae or on stones,
Range: Port Jackson, N.SW. to Perth, W.A.; and
south eastern Tas.
Remarks: This species was known for many years
from the two type specimens (Kaas & Van Belle
1985), which had never been compared together.
Ashby (1921) distinguished his new subspecies
Choriplax grayi pattisoni from the nominate
subspecies by its greater width and its
proportionately smaller tegmentum. He elevated his
subspecies to specific rank (Ashby 1924a) after
examining the holotype of C. grayi in the British
Museum, but did not actually compare the two
holotypes. Kaas & Van Belle (1985) synonymised the
two taxa, which is in agreement with my findings.
Kaas & Van Belle (1985), who have not examined
the holotype of C. grayi pattisoni, state that the
proportionately smaller tegmentum of this
specimen is an artifact of Carpenter’s (in Pilsbry
1892) erroneous measurements of the holotype of
C. grayi. However, they appear to misunderstand
the proportions involved, as they state that the
tegmentum of the holotype of C. grayi pattisoni is
nearly 14% larger than that of the holotype of C.
grayi, but ignore the fact that the former specimen
is 50% larger than the latter, so that the ratio of
tegmentum to total size is in fact smaller. The
holotype of C. grayi (BMNH 1877.11.7.2) is a
juvenile shell with a proportionately larger
tegmentum than the holotype of C. grayi pattisoni
(SAM_ D15019), which is an adult specimen. After
examining the series of specimens available, it is
apparent that the tegmentum is produced at an early
stage of growth and then does not increase in size.
The articulamentum continues to increase in size,
so that the tegmentum in an adult shell (Fig. 1A)
is proportionately much smaller than in a juvenile
(Fig. 1B), and is more central on the valves.
The habitat of this species has been in doubt for
many years. The first specimen found (BMNH
1877.11.7.2) was collected on a stone (Angas 1867),
but of the other eight specimens known, two were
found washed ashore with kelp (SAM DI1S5019,
D17443), one was found on sponge on brown algae
by a diver (AM C151131) and five were found on
red algae by divers (SAM D16542-3;
NMV F51767). The last five specimens were all
pink in colour, matching the host plant, when
collected. The host plant was identified for three
specimens (SAM D16542-3) as the red alga
Sonderopelta coriacea. The identity of the host
plant of two of the W.A. specimens (NMV_ F51767)
is unknown, and could not be determined from the
preserved sample. Previously, C. grayi had been
recorded as epiphytic on kelp (Ashby 1921; Cotton
& Godfrey 1940), in particular on Macrocystis
porifera (Cotton 1964). This arose because the
holotype of C. grayi pattisoni was found washed
ashore with this kelp, and because the colour of the
K. L. GOWLETT-HOLMES
Fig. 2, Choriplax grayi (SAM D16542). A. dorsal view of spirit specimen, 3.5; B. gills, x4.8; C. radula, scale
bar = 10 xm; D. major lateral tooth cusp, scale bar = 10 pm.
CHORIPLACINA (POLYPLACOCOPHORA] 1
dried girdle was very similar to that of the kelp, the
above authors assumed this to be the host plant.
The records from Tas. and W,A, are notable ringe
extensions for the species, and new records for the
faunas of those states.
Discussion
The family Afossochitonidae Ashby, 1925 was
defined by Van Belle (1975, 1981, 1983), Ferreira
(1981) and Kaas & Van Belle (1985) as containing
the geneta Afossochiton Ashby, 1925 (Miocene,
Pliocene), Limrehiton Ashby & Cotton, 1939
(Pliocene), Chariplax (Recent) and Giyprochiton De
Koninck, 1883 (Carboniferous). The distinguishing
feature of the family as defined by these authors
is the presence of unslit insertion plates in all valves.
The type specimens and any other material in
8AM collections of all species of Afossochiton and
Lirachiton were examined to determine the position
of these species and their relationship to Choriplax
grayt, All of these specimens are incomplete, and
Many are in a poor state of preservation,
The members of the genus Afossochiton have
tegmental form and sculpture identical with the
genus Avcanthochitone (Acanthovhitonidae:
Acanthochitonina), fyi are distinguished by raised
ridges on the well developed insertion plates where
grooves and slits would be on the latter genus.
Ajfassochiton is defined as lacking inserGion plate
slits, bur the worn and incomplete nature of the
insertion plates makes it impossible to confirm the
Presence or absence of slits. [ regard A/ossochiton
a5 a gewus within the Family Acanthochitonidae,
aid Alessochtlonidae as a synonym of this family.
Mare completespecimens of Afossochiron with well
preserved insertion plates are required to verify the
position of this genus.
The single species in Lirechiton, L, inexpectus
(Ashby & Cotton 1939) appears to be very closely
related to the extanc species Natoplex (Basset hullit)
matthews! (Bednall & Pilsbry, (894)
{Acanthochitonidae), a relationship also recognised
hy Coton & Godfrey (1940) and Cotton & Weeding
(1941), Lirachiton ts defined as tacking insertion
plate slits, bub I believe the slits are not visible on
Whe available specimens due to the extremely worn
and incomplete state of thetr insertion plates, As
i. inexpectus has the same combinanon of
Pustulose sculpture and grooves as the members of
the subgenus Bassethuiltia, | regard Litachiron as
# junior synonym of this subgenus.
In Choriplax greyi, the structure of the valves and
girdle, and the position of the gills, demonstrate thar
i a unique extant species with characters differing
fron all existing suborders recognized hy Kaas &
Var Belle (1980, 1985) and Van Belle (1981. 19837.
The large number of holobranchial and abanal
gilts preclude its inclusion in the suborder
Lepidopleurina, characterised by few posterior gills
only (Smith 1960; George & George 1979), and the
sombination of characters — large, waslit inseriion
plates, reduced tegmentum, holohranchial gills —
preclude ils assignment co any of the recognized
suborders within the order Neolaciuzta. Recognition
of the family Choriplacidae Ashby, 1928 and the
suborder Choriplacina Starobogatovy & Sirenka,
1975 ts therefore justifled.
Starobogatuv & Sirenko (1975) and Sirenko &
Starobogatoy (1977) regard the Choriplacina as a
suborder of the Lepidopleurida in the subclass
Neoloricata. Their classification includes many
orders and suborders, the majority of which are
poorly defined. In view of the poor definttion and
ambiguity of many of the taxa of these authors, [
prefer to follow the higher classification of Van
Belle (1983), which is well defined and consistent
within the characters used,
Choriplacina is here regarded as a fourth
suborder of the order Neoloricala, because its
combination of characters place it within this order
but distinguish it from the three subartlets
recognized by Van Belle (F983). | believe that the
gills and girdle, as well-as the insertion plates, are
important characters in the higher systematics of
the Polyplacophora. If the gills and girdle are
ignored as distinguishing characters in the higher
systematics of chirons, as proposed by Van Belle
(1983) and Kaas & Van Belle (1985), then the
remaining characteristics of the valves do not
appear to be sufficient to justify the division of the
orders of the Polyplacophora into suborders,
Regarding Glyptochiton, no specimens were
available for study. however, based on descriptions
and illustrations of the genus and species (Kirkby
& Young 1867; De Koninek 1883; Van Belle 1983),
it would scem to be sufficiently distinctive to
warranl recognition of the family Glyplochitonicdae
Starobogutoy & Sirenko, 1975. This family appears
to be related to rhe Choriplacidae, because of its
large. unslir insertion plates and much reduced
leginentum. A detailed examination of
Glyprochiton ts required to determine its true place.
Acknowledgments
I am grateful to Mr and Mrs T. Young who
collected the third known specimen of Choriplex
erayi, which precipitated this investigation, J wish
to thank Ms & Morris (BMINR), Mr L. Lock (AM)
and Ms S3..Bayd (NMY) for the loan of specimens
from their respective institudions, Mr W- Zeidler,
Mr §. A. Parker and Dr W. F Ponder kindly
reviewed earlier versions of the manuscript. Dr P.
Kaas and an anyonymons referee are thanked for
1h K. L. GOWLETT-HOLMES
critical comments. The photographs were tuken by
Mrs.J, Forrest, Ms. A. Renfrey (SEM of radula) and
Mr N. Holmes (BMNH 1877,11.7.2).
References
Apams, H. & Ancas, G. F, (1864) Descriptions of new
genera and species of Chitonidac from the Australian
seas, in the collection af George French Angas, Prac.
Zool, Sac. Lond. for 1864, 192.194,
Astias, G. P. (1865) Descriptions of ten new species of
shells, chiefly trom the Australian seas. Proc. Zool See.
Jiond. for 1865, 55-58, pl. 2.
—— (1867) A list of marine MoUusca found in Port
Jackson Harbour, New South Wales, and on adjacent
coasts, With notes on their habitats, etc. Part [. Zhid.
for 1867. 185-233.
AsHay, E, (1918) Notes on South Australian
Polyplacophora, with additions 10 the fauna; together
with a list of Australian Polyplacophora, showing their
distribution in the Australian states. Trans, R. Soc, §,
Aust, 42, 79-87,
—— (1921) The rediscayery of Choriplax (= Microplax)
grayi, Adams and Angas {Order Polyplacophora), with
notes on its truc place in the natural system) and the
description of a new sub-species. [hid, 45, 136-142.
—— (1924a) Notes on the types of Australian
Polyplacophora in the British Musuem. Mid 48,
328-333.
— (1924b) The regional disiribution of Australian
chitons (Polyplacophors). Aust, Assoc, Adv, Sei 17,
366-393.
Corton, B.C. (19694) “South Australian Mollusca:
Chitons.” (S. Aust, Govt. Printer, Adetaide,)
— & Gonrrey, EK, (1940) “The Molluses of South
Australia, Part UH, Scaphopoda, Cephalopoda,
Aplacophora and Crepipoda.” (8. Aust. Gavt. Princes,
Adelaide.)
—- & Weeping, B. J, (1939) Plindersian Loricates.
Trans, R. Soe. §. Aust. 63(2), 180-199, pl. 7.
—— & —— (1941) The correlation of recent and fossil
Crepipoda (Mollusca) of the Australian sub-region, Rec.
S. Aust. Mus. 4(4), 435-450.
De Kontncr, L.-G. (1883) Faune du Caleaire Carbonifere
de ly Relgique, Quatnéeme Partic. Gasteropodes (spite
afin). dnn, Mus. R, Bist, Nar, Belg, 8, 1-240, pls 12-94.
FERRFIRA, A. J. (1981) Laminoplax, a new gents of
chitons and the taxonomic position of Hanleva dalli
Kaas, 1957 (Polyplacophuora, A fossochitonidye).
Nautilus 95(4), 189-193.
Geoxce, J. D. & Groce, J. J. (1979) “Marine Life, An
Illustrated Encyclopedia of (nvertebrates in the Sea.”
(Rigby, Adelaide)
IkepALE, T. & Huu, A. F. B. (1925) A Monograph of
the Australian Loricates (Phylum Mollusca-Order
Loricaia). Part V. Ausé. Zeal 4(2), 75-L11, pls 942.
— & (1927) “A Monograph of Australian Loricates
(Phylum Mollusea-Order Loricata).” (R, Soc, N.SW.,
Sydney.)
Kaas, PB. & VAN BRILe, R. A. (1980) “Catalogue ol
Living Chitony (Mollusca: Polyplacophors).” (W.
Backhuys, Rotterdam.)
— & (1985) “Monograph at Living, Chitons
(Matlusca: Polyplucophora) Vol. 1, Order Neoloricuta:
Lepidopleurina.” (fi. J. Brill/W. Backhuys, Leidew,)
Korky, J. Wo & Youna, J. (1867) Notes onsome remains
of Chiton and Chiftenellus fram the Carboniferous
strata of Yorkshire and the wesi of Sealand, Geol, Mae,
4(38), 340-343, pl. 16,
KormFeur, A, & WAnscHer, J. H. (1978) “Methuen
Handbook of Colour, 3rd Kdition.” (Kyre Methweti,
London)
Prispry, H- A_ (1892) Polyplacaphora. Jn G, W. Tryon
(Ed.), “Manual of Conchology” Vol, 14, pp. i-wsaiv,
1-128, pls 1-30.
—— (1894) Notices of new chitons IIL. Nuntifus 712),
148-139.
SIRENKO, B. lt. & Staropocatov, Y, 1, (1977) Gn the
systematics of Palgozoic and Mesozoic chitons, Paleent,
Zh, 1977(3), 30-41; English trans!., (1977) Paleons, Jf
T4(3), 285-294,
Smitu, A, G, (1960) Amphineura. pp. 47 76. In Moare,
PC. (ed.), “Treatise on Invertebrate Paleontology. Part
1, Mollusca {" (Univ. of Kansas, Lawrence.)
STAROBOGATOV, Y, 1. & SrReENKO, B. I, (1975) Gn the
systemanics of the Polyplacophora, Jv Likharev, 1. M-
(Ed.), “Molluscs, Their systematics, evolution and
significance,” Lenigrad, Vol, 5, pp, 2-23; English tranal.,
(1978) Malac. Rev, Li(1-2), 73-74.
Van Beure, R. A. (1975) Sur ta classification des
Polyplacophora: UW. Classification systetiatique des
Lepidopleutina (Neolorivala) avec Ja description des
Helminthochi\oninae, nov, subfam. (Lepidopleuridac)
el de Mesochiton noy, gen. (Helminthochitoninae), Inf
Soc, Belge Molac. 4(6), 133-145, 3 pls.
— (1981) “Catalogue of Fossil Chitons (Molluscs:
Polyplacophora).” (W. Backhuys, Rotterdam,)
—— (1983) The Systematic Classification of the Chitons
(Mollusca: Polyptacophora), Inf Soc. Belge Malac,
T4U(1-3), 1-178, pls 1-13.
Zemier, W. & Gow.err, K. 1, (1986) Molluse type
specimens in the South Australian Museum, 3.
Polyplacophora. Ree. S. Aust Mus, 19(8), 97-115.
A NEW SPECIES OF ACANTHOCHITONA
(MOLLUSCA: POLYPLACOPHORA:ACANTHOCHITONIDAE),
FROM SOUTH AUSTRALIA
BY K. L. GOWLETT-HOLMES & W. ZEIDLER*
Summary
A new species, Acanthochitona saundersi sp. noy., is described from Nuyts Archipelago, Spencer
Gulf and Yorke Peninsula, South Australia. The new species most closely resembles A. bednalli but
is distinguished from it by a narrower, deeply grooved jugum and by the spicule arrangement of the
sutural tufts. The new species was located on rocks embedded in sand pockets amongst reef by
divers.
KEY WORDS: Chiton, Polyplacophora, Acanthochitonidae, South Australia, Acanthochitona, new
species.
A NEW SPECIES OF ACANTHOCHITONA (MOLLUSCA: POLYPLACOPHORA:
ACANTHOCHITONIDAE), FROM SOUTH AUSTRALIA
by K. L. GOWLETT-HOLMES & W. ZEIDLER*
Summary
GowLett-Hotmes, K. L. & ZEIDLER, W. (1987) A new species of Acanthochitona (Mollusca:
Polyplacophora: Acanthochitonidae), from South Australia. Trans. R. Soc. S. Aust, 111(2), 111-114, 29
May, 1987.
A new species, Acanthochitona saundersi sp. nov., is described from Nuyts Archipelago, Spencer Gulf
and Yorke Peninsula, South Australia. The new species most closely resembles A. bednalli but is distinguished
from it by a narrower, deeply grooved jugum and by the spicule arrangement of the sutural tufts. The new
species was located on rocks embedded in sand pockets amongst reef by divers.
Key Worps; Chiton, Polyplacophora, Acanthochitonidae, South Australia, Acanthochitona, new
species.
Introduction
The chiton fauna of South Australia is relatively
well known due to the past work of E. Ashby,
W. G. Torr and others (Zeidler & Gowlett 1986).
The last species description from South Australia
was that of Weeding (1940), During studies on the
Acanthochitonidae, one of us (KLG-H) located
what was thought to be an aberrant form of
Acanthochitona bednalli (Pilsbry, 1894) Later
discovery of four more specimens with the same
characters and from similar habitats has led us to
recognise the form as undescribed. Here we describe
this new species.
Materials and methods
The material reported here is desposited in the
South Australian Museum, Adelaide (SAM) and
was collected by “Hookah” diving. All wet material
is preserved in 2% formaldehyde/propylene glycol
solution, Measurements of specimens are linear
when the specimen is laid flat rather than along the
curved surface. The radula was removed from one
of the paratypes (SAM D16698) for examination
after the method of Ponder & Yoo (1976). As the
holotype has not been disarticulated, diagnostic
features of the articulamentum and radula are
described from the above paratype. Colour
descriptions follow Kornerup & Wanscher (1978).
Acanthochitona saundersi sp. nov.
FIGS 1, 2
Holotype: SAM D16699, dry, complete specimen
11.95 = 5.15 mm, collected on edge of granite slope,
under sand in 8 m depth, in cove off NW point of
East Franklin Island, Nuyts Archipelago, S. Aust.
by K. L. Gowlett, 20.vii.1983.
* South Australian Museum, North Terrace, Adelaide, S.
Aust. 5000.
Paratypes: SAM D16698, one specimen,
disarticulated, in spirit 10.9 x 6.15 mm, collected
on granite ledge, under sand at 6 m depth, on inside
of reef off SW side of East Franklin Island, Nuyts
Archipelago, S. Aust. by K. L. Gowlett, 18.vii.1983:
SAM DI17441, two dry complete specimens 12.8 x
6.8 mm and 8.05 x 5.4 mm respectively, collected
on smooth rock, under sand at 12 m depth, on Far
West Bottom, Tiparra Reef, Spencer Gulf, S. Aust.
by K. L. Gowlett, 13.v.1982. SAM D17475, one
complete specimen, in spirit 11.85 x 6.9 mm,
collected on granite fragment in sand pocket on reef
at 7 m depth, off Point Gilbert, Port Moorowie,
Waterloo Bay, Yorke Peninsula, S. Aust. by N. J. C.
Holmes, 29.iii.1986.
Diagnosis: Small chiton to 15 mm. Carinated.
Tegmentum white to cream speckled with brownish
yellow; jugum about 1/6 width of specimen with few
nodulose ribs with deep narrow interspaces; beak
of 4th valve with dark brown spot; pustules “U”
shaped. Articulamentum white, slit formula 5/1/3.
Girdle with alternating bands of white and brownish
yellow; spiculose with prominent sutural tufts of
various sized white spicules.
Description of Holotype: Anterior valve with five
weak ribs, sculptured with distinct, random, “U”
shaped pustules, smaller, but not coalescing near
apex.
Median valves beaked, jugum narrow, 1/6 to 1/3
width of tegmentum, with longitudinal nodulose
ribs containing deep, narrow interspaces, pustulose
posteriorly. Lateropleural areas with longitudinal
rows of distinct, elongated, triangular pustules near
jugum; pustules becoming random and “U” shaped
toward edges. Third median valve with distinct dark
brown spot on beak.
Posterior valve tegmentum slightly longer than
wide; jugum pustulose; antemucronal area with
radially arranged, distinct, triangular to “U” shaped
pustules; mucro dark brown, granulose, central;
112 K. L. GOWLETT-HOLMES & W. ZEIDLER
. .
>
pis
—
Fig. | Acanthachitona saundersi sp. nov. A. holotype, x 10,5 (SAM D16699); B. girdle and sutural tuft, paratype,
«25 (SAM D17441); C. radula, paratype, scale bar =20 ~m (SAM D16698).
postmucronal area slightly concave but not steep,
with distinct, random, “U” shaped pustules.
Girdle with numerous coloured, blunt-tipped
spicules (75-100 «m long, 5 »m wide), and sparse,
long, clear, sharp-tipped spicules 4- 5 times longer,
more numerous toward outer edge. Some spicules
in girdle encroachment at valve sutures and toward
outer edge are clear, blunt-tipped, 3-5 times thicker
and 2-4 times longer than coloured spicules. Sutural
tufts (Fig. 1A, 1B) prominent, with many clear,
tapering, sharp-tipped spicules (100-350 zm long,
20-30 um wide at base) with a few to 2 mm long,
600 xm wide at base.
Gills merobranchial, adanal, with 11 ctenidia on
right side and 10 ctenidia on left side.
Etymology; Named for Mr Frank L. Saunders
(1887-1982) who, as an amateur collector,
contributed significantly to the study of the chiton
fauna of southern Australia and was directly
responsible for one of us (KLG-H) developing an
interest in chitons.
Additional Characters from Paratype:
Articulamentum white; anterior valve slits 5, short,
1/6-1/5 width of articulamentum, in short grooves
1/4-2/3 width of articulamentum (Fig. 2A); median
valve slit 1, short, 1/5-1/3 width of articulamentum,
BREW SOUTH AUSTRALIAN CHITON 3
in shallow groove to edge of regmentum (Fig. 2B);
posterior yalve slits 3, 1/3-1/2 width of
articulamentum, in grooves to edge of tegmentum,
articulamentuim edge between Leeth pectinate (Fig,
2C, ZE)..
Radula (Fig. 1C) with small, rectangular, concave,
central teeth, slightly narrower apically, heads weak;
first lateral teeth narrow, elongate, folded around
base of central tecth: major lateral teeth elongate,
narrower basally, wilh wider tricuspidate heads,
central cusp slightly longer,
Variation: Apart from slight colour variations, the
four paratypes are like the holotype and vary only
in the number of gills, with 12-14 ctenidia om right
side and 11-13 ctenidia on left side,
Range; Central to western South Australia.
Habjiat: Singly on rock embedded in fine to
medium sands, in sand pockets on reefs in areas
usually exposed to moderate swell.
Comparison with other species: 4. saunders} was
compared with ather species. of Acanthochitena in
the collections of SAM, the Australian Museum,
Sydney (AM), the Museum of Victoria, Melbourne:
(NMV), the Western Australian Museum, Perth
(WAM) and the Tasmanian Museum iand Art
Gallery, Hobart (TM). It most closely resembles 4.
bednatli{Pilsbry, 1894) and superficially resembles
A, granostriata (Pilsbry, 1894). Both A. saundersi
and A, hednalli are easily distinguished from other
southern Australian species, including 4,
granostriata, by the jugum which is ribbed in these
(wo spevies but is smooth or pustulose in the other
dig. 2. Aéanthochitona saunders’ sp. nov. paralype (SAM 016698). A. anterior valve; B. median vaive, C. posterior
valve; D, median valye (anterior profile): E. posterior valve (side profile); scale bar =
i mm.
TARLE 1. Distinguishing characters of Acanthochitous bednalli.and A. saundersi SP. HON,
Character A. hednatli
Jugum
Pustules oo valves
Sutural oofts
elongate, drop or oval-shaped
spicules, all of simitar length
Colour white and dark brown; beak of 4th valve
without spor
Habitat intertidal, under rocks in silty areas
width approx. 1/3 width of specimen (dry);
with many ribs with wide inlerspaces
with very numerous, thin, sharp-tipped
A. saunders?
width approx. 1/6 width of specimen (dev);
wath few nodulose ribs with deep narrow
interspaces
“U" shaped
with fewer, slighuly thieker spicules of
various sizes
while and brownish yellow; beak of Silt
valve wilh dark brown spot
sUb-littoral, on rock under sand, in-sand
pockets amongs! reel
114 K. L, GOWLETT-HOLMES & W. ZEIDLER
species. We consider A. johnstoni (Ashby, 1923)
(holotype, SAM D12185) to be a synonym of A.
bednaili, (Zeidler & Gowlett 1986), A. saundersi is
best distinguished from A. bednailli by the
characters given in Table 1, but is otherwise very
similar,
Remarks: The kind of habitat occupied by A.
saundersi and its relatively small size probably
accounts for it not having been collected previously,
It is probably rare as only five specimens have been
collected despite extensive collection of chitons by
SAM divers or associates for the past decade. None
were found in the collections of other Australian
museums.
Acknowledgments
We thank Ms M. A, Garback, Academy of
Natural Sciences of Philadelphia, U.S.A., for the
loan of the syntypes of A. bednalli (ANSP 64955)
and A, granostriata (ANSP 64847) and the
following curators and collection managers for
providing data and specimens from collections held
by their respective institutions: Mr I. Lock (AM),
Ms 8S. Boyd (NMV), Dr F. E. Wells (WAM) and Ms
E. Turner (TM). The photographs were taken by
Mrs J. Forrest and Ms A. Renfrey (SEM). Mr Nigel
Holmes is thanked for his many collecting efforts
resulting in the discovery of the fifth specimen of
A. saundersi.
References
Kornerup, A. & Wanscuer, J, IL. (1978) “Methuen
Handbook at Colour, 3rd Edition.” (Eyre Methuen,
London.)
Ponper, W. F. & Yoo, E. K. (1976) A revision of the
Australian and tropical Indo-Pacific Tertiary and Recent
species of Pisinna (=Estea) (Mollusca: Gastropoda:
Rissoidae). Rec, Aust, Mus, 30(10), 150-247.
WEEDING, B. J. (1940) A new Flindersian chiton
Acutoplax cottoni. Trans. R. Soc. S. Aust. 64(1), 48-49,
ZEIDLER, W. & Gow eTT, K. L. (1986) Molluse type
specimens in the South Australian Museum 3. Poly-
placophora. Rec. 8. Aust, Mus. 19(8), 97-115,
REIDENTIFICATION OF CTENOTUS SCHOMBURGKII (PETERS, 1864)
(LACERTILIA: SCINCIDAE)
Summary
BRIEF COMMUNICATION
REIDENTIFICATION OF CTENOTUS SCHOMBURGKII (PETERS, 1864)
(LACERTILIA: SCINCIDAE)
Lygosoma schomburgkii Peters! was described in 1864
from a collection of reptiles made by Richard Schomburgk
of Buchsfeld, South Australia (= Loos, 4.5 km W of
Gawler2). Peters neither indicated the number of
specimens on which he based the species, nor whether all
of the collection was from the vicinity of Buchsfeld.
However, with the exception of Moloch horridus (“ein
einziges verstiimmeltes Exemplar”), a single specimen of
Litoria adelaidensis} and Ctenotus schomburgkii
(auctorum), all of the 42 species of reptiles and amphibians
recorded in the paper (on modern synonymies) have been
subsequently recorded within 50 km of Buchsfeld (our
data: T. D, Schwaner, pers. comm.).
After devoting “a month to the examination of the
specimens in the Berlin Museum”, at which Peters was
based, Boulenger4 placed L. schomburgkii in the
synonymy of L. /esueurii, diagnosed in the generic key
as having 24-32 midbody scales, feebly keeled subdigital
lamellae, four supraoculars and prefrontals in contact,
However, C. schomburgkii as currently recognised has
mucronate subdigital lamellae and prefrontals usually not
in contact5®,
Storr> examined four specimens identified as types of
Lygosoma schomburgkii in the collection of the
Zoologisches Museum, Universitat Humboldt, Berlin
(ZMB). Two species were represented and ZMB 4713a
(SVL 45 mm) was designated lectotype® (not Storr® as
listed by Cogger ef ai.7), One of the paralectotypes (ZMB
4713b) is conspecific with the lectotype, while the others
(ZMB 4713c-d) were subsequently® (as ZMB 4719c-d)
included in the paratype series of Ctenotus uber orientalis
Storr®. These latter two specimens were re-registered
ZMB 41236-37.
Subsequent collections of reptiles from about the type
locality by us, and examination of South Australian
Museum specimen records, have revealed only C. uber
orientalis (AM R59944, Thompsons Beach; R115921, 6.3
km S Mongalata; R115926, “Tracy”, 4.8 km N Mongalata;
R115979, Middle Beach; R115989, R115991, nr “Glenview”,
Barossa Range; SAM R11202, 1.6 km N Tea Tree Gully;
R15034a-b, Pt Prime; R15141, Sturt Ck, Adelaide Hills;
R15617, Golden Grove; R16543, Morialta Conservation
Park; R16867, Belair Recreation Park) and C. robustus
Storr (AM R111491-92, 4.5 km SSE Gawler; R115878-79,
Burra Creek Gorge; SAM R1703, Bute; .R2093, Fifth Ck,
Montacute; R8144, Hummock Mt; R9983-86, R9992,
R9994-95, Waterfall Gully, R10006, foot of Mt Osmond;
R11208, Eden Hills, nr Sturt Ck; R11209, 1.6 km N Tea
Tree Gully; R14192a-b, Aroona Rd, Reynella; R27413-14,
4km W Palmer) from the Adelaide Plains and Mt Lofty
Ranges. The latter is presumably Peters”! “12. Lygosoma
australe Gray sp. = L, Lesueurii D.B.” There have been
no subsequent records of C. schomburgkii (sensu Storr
and subsequent authors) from the Adelaide Plains, the
nearest record being from Bungunnia Station (33°50'S
139°50'E; South Australian Museum R15045), over 120
km from Buchsfeld, in the Murray Mallee.
A translation of the type description of Lygosoma
schomburgkii reads:
“Stands between L. /abillardieri and australe. Head
and tail brown, back black, with four white or green
stripes, of which the two middle ones are the thinnest
and are almost twice as far apart from the outer stripes
as from each other; on either side of the body is a line,
which is twice as far from the outer back stripes as they
are from the middle back stripes which lie next to them;
between each outer and inner back stripe is a line of
white spots; between the side stripes and the outer back
stripes are three to four rows of small white spots, and
below the side stripes a few largish speckles, which often
blend in with the greenish-white belly.”
In the nomenclature of subsequent workers on
Ctenotus, Lygosoma schomburgkii has a pair of
paravertebral stripes (Peters’ “middle back stripes”), a
single laterodorsal line of white spots between the
paravertebral and dorsolateral stripes (“outer back stripes”)
and three to four lines of small spots in the upper lateral
zone between dorsolateral and midlateral stripes (“side
stripes”) on a black dorsal ground, Our interpretation is
supported by the stated degree of separation between
stripes.
South Australian populations of Cfenotus
schomburgkii (auctorum) are of the eastern colour type
of Storr5 (Fig. 1A), and have an irregular orange
laterodorsal stripe between an orange paravertebral and
a white dorsolateral stripe (i-e., six light dorsal stripes),
and a single row of large orange squarish to oblong
blotches between the dorsolateral and midlateral stripes.
Eastern populations do not have a single laterodorsal row
of white spots (this being replaced by an orange stripe),
and no population of this species has three to four series
of small spots between the dorsolateral and midlateral
stripes. Specimens ZMB 47l3a-b (Fig. 2A) are
representative of this species. Both have the prefrontals
separated.
In contrast, the populations of C. uber orientalis from
about Buchsfeld (Fig. 1B), including ZMB 41236-37, agree
in all respects with the description of Lygosoma
schomburgkii Peters, although the white midlateral stripe
may be absent in some individuals and the dorsal ground
is more brown than black in some (but not all) specimens
from coastal situations. A white midlateral stripe is present
in ZMB 41236-37 (Fig. 2B).
As Peters did not designate types in his description, and
as the four specimens formerly included in ZMB 4713 were
untagged, and probably registered after the publication
of the description (R. Giinther pers. comm.), we consider
that the type status of the two syntypes ZMB 4713a-b is
open to question, and as these specimens do not agree
with the type description or Boulenger’s? subsequent
definition, and are representative of a species not known
from near the type locality, we reject their type status. If
ZMB 4713a is not a syntype, Storr’s> designation of this
specimen as lectotype is invalid.
The Zoologisches Museum collection contains frogs
purportedly collected by Schomburgk from Adelaide?
additional to those listed by Peters!, suggesting that
Schomburgk sent material from South Australia to Peters
subsequent to 1864 and supporting the possibility that
Fig. 1. A, Crenotus schomburgkii auctorum (Australian Museum R114482; Lake Mungo National Park, NSW);
B, Ctenotus uber orientalis (Australian Museum R115926; 4.8 km N “Mongalata” HS, SA),
ZMB 4713a-b were sent to Peters after the description of
L. schomburgkit.
We consider that ZMB 41236-37 are syntypes of L.
schomburgkii Peters, and we designate ZMB 41236 as
lectotype.
This reassignment of the name Lygosomta schomburekii
necessitates four nomenclatural changes. C. wher orientalis
Storr® becomes a synonym of C schomburgkii (Peters),
while C, wber uber Storrs and C. uber johnstonei Storr’
become C. schomburgkii uber and C. schomburgkii
johnstonei respectively. The only available name for C.
schomburgkii (auctorum) is Lygosoma fiseheri
Boulenger**:7, a nomen novum for Hinulia muelleri
Fischer?, at that time a junior secondary homonym of
Scincus muelleri Schlegel’, Although the types of H.
muelleri cannot be found’, the description and figures?
clearly apply to the western colour type of C
schomburgkii (auctorum),
H. G. Cogger kindly allowed access to unpublished
notes and reproduction of photographs of the types
W7
Fig, 2. Syntypes (sensu Storr) of Lygosoma schomburgkii Peters. A, ZMB 4713a (upper; lectotype of Storr)
and 4713b (lower) and B, ZMB 41236 (upper; lectotype designated herein) and 41237 (lower; paralectotype).
(Fig. 2), and provided useful comments on the manuscript.
T. D. Schwaner and A. Edwards permitted us to examine
South Australian Museum material, and checked locality
data, N. Shea translated much of Peters’ paper. B, Coulson
provided Fig. 1B. A. E. Greer and R. Sadlier also
'Peters, W. (1864) Mber. K. Preuss. Akad. Wiss. Berlin
1863, 228-236,
2Praite, R. & Tolley, J. C. (1970) “Place Names of South
Australia.” (Rigby, Adelaide).
3Tyler, M. J.(1985) Mitt. zool. Mus. Berl. 61, 335-337.
4Boulenger, G. A, (1887) “Catalogue of the Lizards in
the British Museum (Natural History). 3.” (British
Museum, London).
5Storr, G. M. (1969) J. Proc. R. Soc. W. Aust. 51, 97-109,
commented on the manuscript.
Our field work was supported by a grant from the
Reserves Board of South Australia. B, Coulson and B.
Miller are thanked for field assistance,
6 (1971) Rec. S. Aust. Mus. 16(6), 1-15.
7Cogger, H. G., Cameron, E. E. & Cogger, H, M. (1983)
“Zoological Catalogue of Australia. 1, Amphibia and
Reptilia.” (Aust. Govt Publishing Service, Canberra).
8Storr, G. M. (1980) Rec. West. Aust. Mus. 8, 441-447.
Fischer, J. G. (1882) Arch. Naturgesch. 48, 281-302.
Schlegel, H. (1837) “Abbildungen neuer oder
unvollstandig bekannter Amphibien, nach der Natur
oder dem Leben entwofen.” (Arnz, Disseldorf).
M. PETERSON, 69 Alvah St, St James, W.A. 6102 and G, M, SHEA, Dept of Veterinary Anatomy, University of
Sydney, N.SW. 2006,
ON THE INDETIFICATION OF CTENOTUS SCHOMBURGKII (PETERS)
Summary
BRIEF COMMUNICATION
ON THE IDENTIFICATION OF CTENOTUS SCHOMBURGKI! (PETERS)
In preparation for a paper on the Ctenotus of the
Eastern Division of Western Australia, I borrowed the
types of Lygosoma schomburgkii Peters from the Berlin
Museum. Two of the specimens belonged to a wide-
ranging Western Australian species; the other two could
not be matched with any Crenorus | then knew (they were
later found to agree with C. uber orientalis Storr, 1971),
I therefore settled the name schomburgkii, by lectotype
designation, on the taxon I knew.
Twenty years later, when my usage of the names
schomburgkii and wber is well-established in the literature,
Peterson and Shea attempt to upset the above designation
on the grounds that the lectotype and paralectotype were
not part of Peters’ original material. They argue thus:
(1) Legosama schomburgkii (sensu Storr) does not occur
within 50 km of Buchsfeld;
(2) Boulenger, alter examining specimens in the Berlin
Museum, placed L, schomtburgkii in the synonymy
of L, lesweurii; and
(3) Peters’ description does not fit Crenotus schomburgkii
(sensu Storr) but agrees “in all respects” with C, uber
orientalis Store.
The first two arguments are frivolous grounds for
upsetting a lectotype designation, and the third is untrue,
There is no evidence that Schomburgk ever provided
Peters with locality data. If he had obtained his specimens
of schomburgkii in the Murray Mallee, would he have
informed Peters of it? The locality “Buchsfeld” applies
more to Schomburgk’s place of residence than to the
provenance of his specimens. Al any rate Peterson and
Shea admit that other specimens, apart from Lygosoma
schomburgkii, could not have come from the vicinity of
Buchsfeld.
Boulenger’s concept of Lygosuma lesueurii was so
grossly composite that it tells us nothing about the
specimens of L. schomburgkii that he may have seen in
Berlin. The subdigital lamellae of L. pantherinum Peters
are finely keeled (as in L. schomburgkii) but that did not
deter Boulenger from synonymizing pantherinum with
lesueurii (a species whose subdigital lamellae are widely
callose),
was composite. His description of the back as “black with
four white or green stripes" could only apply to spirit
specimens of eastern Cienotus schomburgkii (sensu Storr),
in spirit specimens of C, uber orientalis the back is brown
or grey and the stripes brownish white or greyish white.
Peters’ description of the side of the body is almost as
certainly based on the specimens of C. uber orientalis.
It is not surprising that Peters was unable to separate
two such superficially similar taxa as Ctenotus
schamburgkii and C, uber orientalis (no one could until
! pointed the way). Believing his material to comprise a
Single species, Peters was free to base his description on
any or all four of his specimens; and I, as first reviser,
was equally free to choose any of them as lectotype.
G. M. STORR, Western Australian Muscum, Francis Street, Perth, W.A. 6000,
ON THE OCCURRENCE OF BRACHYLAIMA SP. (TREMATODA) IN THE
FERAL HOUSE MOUSE, MUS MUSCULUS, IN SOUTH AUSTRALIA
Summary
TRIER COMMUDICATION
ON THE OCCURRENCE OF BRACHYLAIMA SP. (TREMATODA) IN THE FERAL
HOUSE MOUSE, WUS MUSCULUS, IN SOLTH AUSTRALIA
Durtigs study al the ecology af the house mouse, Mus
measeufus, conducted in the Lower light area of Seurh
Australia, mice were crapped overnight in break-back Lraps.
Inthe moming the stomachs aud intestines were removed
from (he dead mice and preserved in 70% erhanal for eut
anulysis. Trematades identitied as Broctwlittnre sp. were
found inthe small intestine of nine of 9§ mice examined
herween February 1985 und Junge 1986. The greatest
numbers of infected mice were found in September’
October 1985S, Infection data lar ahe period Septeniber
1985-Tune 1984 are more eeliable (han earlier observations,
During ts period, seven of 48 animals were infected. Mhe
resiricuon of the infectian ta the small Intestine is not
Certain as examination of olher parts of the cul was less
thoraugh,
Numbers of irematodes found it individual mice ranged
trom one ta 34. Trematode-infected mice were found in
ull parts of the study area, living Under dense roadside
vegctation from one iw five Kilometres gorth of Lower
Light,
A large aumber of Srchkelaima app. have been
described from the small intestine and cuecum ot birds
and arnumals, but only three species are recounised from
Australia"! Iwo are from selall marsupials and ane
from a bird, Aracéelaie desyuri (Ss. J. Johnston. r91ayt
was dea¢ribed from the nalive cat, Dasvures viverrinus,
“itt the vicinity of Sydney, NSW"
Sandars, (1957) pave a further description ot &,
dasyurt Tram the short-nosed bandicoal, Lseadan
olesulus, from Queensland, and placed A simile (S.J,
tohnston. L93) described from Masvuras v/verrinus, as
asynonym of A daspurt Brochylaime pulehellans (S.J.
Johnston, 1917)" was described from the Wonga pigeati,
Leucusarcie melenmonea, from Nocthwestern NSW, and
Brachvlaimea antechini Peisley & Howell, 19757 was
described from marsupial mice, “afechinus snuartié and
A. swaiasani, trom the vicinity of Canberra, AST.
Because of the method of preserving rite mice with the
(etiatodes sullin the alimentary canals, the tremiaicaes
were rather distorted and wWeatificaion was difficult. Two
speeimens which were recovered from a deep frozen mouse
(2.A1L.P98S] were fixed in formal aleobol under coverslip
pressure. They measure (1) 4.0 mm by 0.5) mm ane (2)
3,9 mm by O.d4 mim, These measurements lie within the
ranges siver for the three Australian Brocfiviaiine spp. We
have only been able to get a comparison of sizes of rhe
oral sucker and thy aeetabulum in one (flattened)
spechmen, fhe diameter of the cral sucker was 0.24 mom
and of the acetabuthim 27 myn.
in deseribing & entechini, Peisley & Howell did rot
menuon & diasevuri (syn. Hermasionnum dusvuri) and it
16 possible iat Cumnparison of the bwo species will show
ia & avteehin’ is a synonym ot KB desveri.
Certainky: Que species appears ro resemble & dus puri
al & anteckia’ mare closely than i does & pulchedturn.
The figures of & ouichellynr differ [ram those of the othey
TWO species HL Ue relationship of the organs": alse,
Jolwistyn states that (he gonails are not us lose tuverher
zs iy usimil in the pends, However, wu species must be
consklersd distinct Mom & desvuri and & oetewhin
bevause of the forward extent of the ierus. The uterine
coils in B dasywri have been described as reacluny the
middie of the agetabulua, and reaching the posterior
matin af the acetabulum. In A entecfeni they are
confined posteriatly to the acerabulum’, In the
speciniens examined here, the uterus extends rom beyond
the anterior burder of the acetabulum (o the region of the
pharynx.
ic is unlikely Chal the species-was introduced with Mus
nixscalus in the early days of aceupation of South
Australia, ‘Tematades have not been recorded previously
in ML enjéseie/us int Austeatia'®, The latler study occurred
in habitat and climate thar are very similar i those at
Lower Light. and the area way subject to similar agri-
cultural practices Filteen species of endo- and ecto-
(purasiles were recorded, but did not include tremaiodes,
Tn a continuation af the Vieronan scudy”, a further 98!
mice colleetad at megular intervals since November 1983
wert dlyo recently examined, and found to be free of
trematodes,
In. 1971"? thirty species of Brachy aint were recognised
aad since then some sixty more have heen deserbed, Four
of the species recorded by Yaraguri!” were from Mf
muscles, B riuseuli (Rud, L819)", Bo recervum Duj.,
1845", A mequans (Loass 1899)" and & peroniysct
Reynolds, 1938!, the last three as synonyms of &
muscu, A dequans and B recwrvem were recovered
experimentally fram white mice © B rudninie Mase
Coma & Monto, 1986'7 was found. naturally and
experimentally, in Aduy musecu/us in. Spain.
Nisalsa untkely that the ematode hay bern acquired
from Rarius ractas, which has been ¢aprured at the study
site, ar R. norvegicus which may have been in the arew
in The past and may be still extant!®, Alihough a few
Brichviaine spp. have been found expert ientally in cats,
only & migrans Dujardin, 1845! Crom Afus ratty (now
Kars rattys) and Mus decwmunus (now Rattus
forewwews) in Europe, and B nerti Bagh, 1962!" (trom
Rattus ratius in Midial are listed a9 natural infections!.
& ishigakiense Kamiya & Machida, 197774, has since
been recorded from R. natives in Japan, and Brachylaima
ruminge Mas-Coma & Montoliu, 19867 from Ro raftus
ia Spain,
Iris mast likely that the tremutode originated from a
sinall mammal (though as. bird cannot be cecluded) which
was nee, or perhaps still is, an nihabirant of the general
area in which che trematade Was found, The animal must
include snails in its food 4s these are second Intermediate
hosts in all seven species of Hracky aie tor which life
histozy data are available”,
Twerry-nine maninials have been recorded within a radius
of 3) kin of 34°32'S, 1L8°20"R (Lower Livtn)!*, The
habitel within this area is nor honivesneous, Three af these
mammals (Teehaswrus vulpecula, Betiongia lesuewr and
Sacewaimes Jlaviventris) are known to be or to have been
in the study area, others may have evry it the area in.the
past Whilt some Wav be still extant, Nine small marsupials
and lau rocdunts are vonsidered as possible original hosts
122
ol our Brachylaina sp. on the basis of dietary habits, The
marsupials are Dasvurus maculatus, Do viverrinus,
Phascogule tapouiusa, Perameles hougainville, Macratis
fonoris, Antechinus flavipes, Sminthopsis murina, S.
crissicaudata aud Isuodon obesubes, and the rodents ire
Hydronys vhresogaster, Rartus fuselpes, R. lutrentus and
Noaromys mitcheltii.
We cannot exclude the native rodents as original hosts
of our Brachyiaimasp., bur we eansider that they are less
Nkely hosts than ure the marsupials. The presence in the
study area al any time of the last three rodents on ow
list iy Cousidered doubiful!®. Aydromyy vhrysopasier
almost certainty 1s or was in the general area and is very
common i newhbouring areas. Brachy/ofnine was not
founcl in 49/7. chrysogaster examined (rom S_ Aust. and
six from Vietoria**, or in 32 from Tasmania’, No
Trematodes were found (0 47 Rattus Seseipey [ror
Vieloria’?. Aruchylaima sp. was collected fram
R. fuscipes trom Brindahella Ranges, A,CT?*, Thirty
& fuscipes fed with metacereariae of Brachplaine
aatechint did not develop adult trematodes, nor did seven
white mice’,
Of the recorded hosts of Brachy aime spp. in Australia,
Desyurus viverrinus and Iroodon obeselus are regarded
as once possible inhabitants of the stucly area, Thoueh
Despyurns vivecrTus 1s now lovally eXtinel, and fyeedorn
obesulus probably extinet in that area, the latter is still
living in the Adelaide Fills, and may have heen formerly
in the Dower Tight areal
Antechinus flavipeois the only species of the genus
which has possibly been in the area. UP muy still be extant.
Antechinus swainsoni and A. stuariti have never heen
found in-S, Aust,
Lencosdrdid Mélangleuce does toy oceur jn Seth
Australis. Native birds evident in the experimental area
' Muckerrax, M.d_ (1498) Prac. Linn. Soe, N.SW. 83,
, 160,
* Spratt, D. M., Beveridge, 1 & Walters, kK.
(Unpublished) “A catalogue of Australasian
monotremes and marsupials and their regarded
' helminth parasites,”
” Mawson, P.M., Angel, L. M. & Edmtoneds, 5, J, 11986)
Ree. S. Aust. Mus. 19, 219-325.
‘ Jotmston, §, J-(1913) Prac, Linn. Soc, N.SAV, 37,
727-740,
" Sandars, 1. F (1957), J. Helminth. VW, 263-272,
8 Serie 8. J. (1917) J. & Prov, R. Soe, NSW. 30,
|
* Poistey, FR. & Howell, M4. (1975) Ini, Jour, Parasil,
, S, 441-447)
9 () buntgleton, G, BR. (1985) Aust. Wildl, Res, 12, 447-445
" Singleton, G. R. (June 1986) Pers. Comm.
'"-Yamaguti, S. (1971) “Synopsis of digencti¢ trematedes
of vertebrates", Vol, 1, 1074 py, (Kkeigaku Publ, Co.,
Tokyo).
N" Rudolph, C A, (819) “Entozoorum synopsis ai
sees mantissa duples ¢t indices looupletissimi ”
ervin,
'F Dulaedin. F. (1845) "Histoive natoeelle clos Helminthes
OU vers intestingaus", Paris,
which are likely to eat snails are Gyrnarhina ribicern,
Corvus coronvides, C. mellori, Colluricincla harmonica,
Strepera versicalor, Corcorux melanorhamphos and
Pomatastomus supereiiiosus, and the introduced birds,
starlings and blackbirds, We know of no trematodes from
starlinws or blackbirds in Australia, and there are no
records of Brachylaima spp. ‘from the native birds
mentioned,
Life History: Metacercariae have been found in the kidneys
of It.of 30 of the gustropod Strangesia capillacea, from
the areas where Aprectinus sruaeri/ was rapped’. There
were no other stages found — ie. nv cercuriae in slime
trails nor sporoeysts in the digestive gland, Two other snail
genera were not infected, Feeding experiments showed that
the metacercuriae were the larval stage of Brachyleima
antechini.
Although we fave looked for snails in the study area
singe we first became interested in che trematodes from
mice, we have Not found any native species, The introduced
snails, Cernuelle virgata and Thebe pisaru, are locally
abundant.
As Tice in the area are known ta eal a yanety of
invertebrates, infection of the mice presumably oceury by
eating snails containing metacercariac, Invertebrate rissue
was found in the stomachs of 39% of mice living in and
adjavent 4) rice stubble im N.S andl it is considered likely
That mice would ent snails when availabte*, Inveriebrare
issue ts als0.a common dlary componenl in mike from
the Victorian mallee’.
We wish to thank friends and colleagues for advice and
help given in this project
" Leoss, A, (1899) Zool, Jahrb. Syst, 12, 521-784.
; * Reynolds, B.D. (1938) J. Parasit. 24, 245-248.
5 Punin, V. ta. & Sumenkava, N. 6 (1963) (Russian text)
Triidy. Inst) Zool Akad. Nauk. Kavakhxk, SSR, 19,
83-8.
'S Timon-David, J. (1954) Campi, Rend. Soc. Biol.,
Paris. 148, TARTU,
i Mee Cpalty, & Montolin, 1. (1986) Z. ParasiiKele. 2,
'* Kemper, C. (1956) Pers, Comm,
Lb * Baugh, SC (1962) 2. ParasiiKde, 21, 502-512,
" Kanelyt, H. & Machida, MM, (1977) Rul. hath. Sei
Mus, Ser, A. (Zool). 3, 125-129,
4 Vamaguti, 5. (1975) “A synoptical review. of life
histories of digenetic trematodes of vertebrates with
special reference to the morphology of their larval
forms,” 596 pp. (Keigaku Publ, Co,, Tokyo),
* Angel, L. M. (1938-1967) Unpublished data,
Obendort, D. L. & Smales, L. RB. (1985) Aust. J. Zool,
33, 33-33-
4 Cyhendor?, 1,4; (1979) Aust, 4. Zool, 27, 867-879,
*5 Spratt, Uk ML (1986) Pets. Comm.
*> Rumford. M. (1985) “Food quality, diet and
reproduction of house mice on irrigated cereal farms.”
Unpobl. PhD. Thesis, Australian National University
ery
=
L MADELINE ANGEL, Souh Austealian Museum, Norih Terrace, South Australia. ane G. J. MUTZE_ Vertebrue
Pests Cantera! Auihority, Depaetment of Agriculture, South Australia.
REPRODUCTION AND GROWTH IN CAPTIVE DEATH ADDERS
ACANTHOPHIS ANTARCTICUS (SQUAMATA: ELAPIDAE)
Summary
BRIEF COMMUNICATION
REPRODUCTION AND GROWTH IN CAPTIVE DEATH ADDERS
ACANTHOPHIS ANTARCTICUS (SQUAMATA; ELAPIDAE)
A captive calony of 4Acunthaphis antarcticus has been
maintained at Whyalla, South Australia since 1975, Several
papers have described the birth of individual elutches into
the colony!?4. Mirtschin has outlined (he maintenance
schedule for the colony’ and the release of death adders
bred at Whyalla’, This paper presents new data on the
biology of A. antarcticus kept at Whyalla, including
seasonality of mating and births, primary sex ratio, clutch
size, caudal luring, growth and maturicy.
Adult snakes were callected from Iran Duke (33° 18'S,
137'08'L), Port Germein (33¢01'S, 138"00'L), Tumby Bay
(34°23'S, 136°06'E) and Ardrossan (34°25'S, 37954'E).
They were housedt in enclosures deseribed by Mirtschin*
ar temperatures: of 27-30°C and ed entirely on house mice
(Mus musculus) and rats (Rattus norvegicus). Neonates
xecepted pink mice, Snakes were fed older, larger mice or
rats commensurate with increasing body size and their
ability to consune larger food items.
Dates of mating and births have been recorded from
the foundation of the colony. Weight (gm) using a Mettler
top-loading balance and total length (cm) using a
centimetre rule were measured for snakes in the colony
of known age in February, June, August and December
1982. Snakes were measured and weighed from clutches
barn in 1979, 1981 and 1982. Primary sex ratios were
determined by probing® for seven cluiches born between
January 1982 and April 1983 (Table J). Chitch size was
défermined for 14 clutches borti between February 1976
and April 1983,
Length and weight against jime and length against weight
were examined by least-squares regression’. Linear (Y =
a + bX), exponential (In¥ = a + bX), logarithmic (Y
a + bInX) and power (InY = a + bln X) equations
Were used to find the besr fit ta ihe data, where Y = the
dependent variable, X = the independent variable, a =
the y-intercept and b = the slope of the fitted fine. For
each regression the coefficient of determination (R2) was
used as the criterion for which equation best fit given
ser of data, Daw were treated without regard to sex,
The A. avturcticus. reported. ‘here show an annual
reproductive cycle’, mating shortly after winter
Tattt t. Dales af birth, sex ratios and clutch sizes for
death adder clutches born in captivity at Whyalla
Clutch Date male female umnsexed total
1 10. 111.1976 — _ — 24
2 20.41.1976 — oe -- 19
a 13i37.1977 — — = 1G
4 10. ii).1979 = = = 24
5 28,i.1981 — oa — IL
6 3su18RP | = = 18
7 G. ii. 1982 7 8 2 i7
8 10,44, 1982 12 Is — 27
9 10.11. 1983 7 17 — 24
10 10,0, 1983 4} 11 — 16
1) Vii, 1983 16 17 _ 33
42 ZL ii, 1983 7 14 _ 21
3 Vi 1983 8 17 — 25
14 221984 ¥ 7 — 16
mi
brumation and giving birth betore brumation the
folluwing year, Mating was observed 24 times prior to
March 1983. and fourteen clutches lotalling 285 neonates
were born into the colony during the same period (Fiz. 1.
Table 1). Mating was at its height (n October and births
peaked in February and March, Gestation takes four-five
montis over summer. Minor mating activity occurs
shortly after parturition.
15
Y
O
Z
a
<x
=
10
i
=
ae
an)
Le
* &
oF
Lu
ag)
=
=,
2
0
JFMAMJJASOND
MONTH
Fig. 1. Monthly ovcurrence of births (open. bars) and
inalings (solid bars) recorded in A. aatareticus at the
Whyalla Tauna Park, South Australia between 1976
and 1983.
Mating has been reported previously for A. antercricus
in January-April!!! and August-December? 0. Shine!?
found that ovulation fakes place in late spring to early
summer and embryos were present in females collected
belween December and March, Males were in
reproductive condition throughout the year,
One female A. entarcticus born in 1980 is known to
have produced clutches in four successive years. Hay &
Magnusson!¢ reported a female who bore clutches For
three years in succession, However, Shine!* found that
approximately 50% of wild adult females were not in
reproductive condition over the summer months and
canchided they show repraductive asynchrony such that
individuals reproduce biannually but the popolation
reproduces each year, Discrepancies between annual
reproductive cycles reported for captive death adders and
biconial cycles in the Field are probably due ia # shift in
Une relative adaptive value of low frequencies of
124
feploduction iit captive versus field conditians!*. Food
availability and low thermal stiess may be prime factors
involved! here,
The meao primary sex ratio differed signiticanny trom
expected’, with females ournumbering males (X2fdr = 1]
~ 44.032, Po0.001), although there was considerable
variation between clutches (Table 1). Hay &
Magnusson!” have-alse noted a significant bias Loward
fernales in death adders. The variation trom oxpected sex
tario cannot be explained by differential reproductive
investments by females to hatchlings of each sex ‘Twenty
seven’ fermale neonates from clutches 7 & 8 (Table 1)
measured within 24 brs of parturition were significantly
heavier (x =5,.84,5D — 0,426¢m, range — $,22-6.65)
thin 15 mates from rhe sare clutches (x = 5.09, 5D -
1,269, tange ~ 4.38-5.54) (t{dl =40] = 6.499, p< 0,001),
Datla contained in Shine’s!? table | show wsignificantly
higher number of adull males than-adult females in wild
populations (N2[d=1] = 4.300, P<0.05). As elapics
hay: heteromorphic sex chromosomes!? if is probable
that sex determination in .4. antarcticus is genetically
controlied'5, [tis interesting to nite that the sex ratia
bias reverses between captive nconates and wild adults,
indicating differential mortality between the sexes in wild
populations.
Clutch sizes of 2-24 have been reported previously for
A. gartarcteus!!?_ The captive clutch sizes reported here
(Table 1) (x = 20.4, SD = 6.31) and by Hay &
Magnusson! (x ~ 20.8, SD = 2.30, N = 12) ag
considerably larger chan those reported for clutches ia
the tleld (x = 7,9)!2.
Neonates have been observed on several occasions
caudal uring immediately pos-parturition, This indicates
that caudal luring is an innate behaviour and Hot learned.
Heatwole & Davison!® found that most species which
exhibit caudal luring do so only as juveniles. Adult dearh
adders in the Whyalla colony have always been observed
10 Whiggte their tails vigorously whenever an enclosure
is approached and especially when a food item is placed
in the enclosure, Carpenter, Murphy & Carpenter also
have reported adult Acanthophis caudal luring!?.
At parturition neonates averaged 15,85 em (SD - L2u)
10 total leagui and $.93 ym (SD = 1.09) in weight. Growth
by weight was best described by the exponential equation:
MW = 1.822 + O,123T
where W = weight (gm) and T = age (mop (Pig. 2).
Correlation coefficients (r) were significantly different
hetween exponential and logarithmic models
(P<20,001)'* but linear aud power models were similar
(uy the exponential model (P's5> 0.05) (Table 2). Growth
by total Jength was best described by rhe linear equation:
L — 15.907 + 1.2U1T
where T is as above, aud L = length (cm) (Fig. 3),
Correlation coefficients of exponential and lowarithimic
models where significantly different from the tinear model
(P's<0.001) but the power model was similar (P >0.5)
(Table 2). The relationship between weight and total
length is given by the power equation:
InW = -6,761 + 3.088InI,
where W and |..are as above. Correlation coefficients for
tle Linear, logarithotic and exponential models were all
500 a
Bi °°
1 r* +
zt
100 So
= * +
mM .
sl ' s+
om , *
oe
& ’ ' x (yet 29+ 128e
Gi ONO) hg Ne zou
2 | mO.97 (Ped Hor)
] .
ee ee a |
Q 12 24 56 44
AGE (mo)
Fig. 2. Growth by weight (gm) of captive 4, aetarcticus
over a four year period, Fach symbol may represent
More (han one data point,
significantly different from the power modet (P's < 0,005)
(Table 2, Fig. 4).
In the field newborn A. antereticus appear in autunin
measuring 42 cm snout-vert length and grow to 20 em
hy the end of che calendar year and 30 em by 24 mol,
This curve considerably underestimates the curve reported
here over that time period for captive animals, but it does
predict actual fourth year size accurately if extended on
ut the same gradient. The results reported here may be
taken as an apcmun curve lor captive death adders. The
initial difference in the growth curves may be because
juvenile death adders at Whyalla are fed mice and wild
populations take mostly reptiles in younger life’.
Wild A. aatarcticus do not reach sexual maturity waril
24 and 42 mo for males and Semales, respectively’.
TABLE 2. Comparison of regression ediations Jor
grow?h on weight, grow/h on total fength and weigh? vn
total length in captive death adders at Whyalla.
Equation R?
Weight vs Ape
W= 4.009 + 9.434T 0.869
Wo 230.233 + 147.134In1 N.O4z
InW R22 = 01236 0,942
uW = 0.217 + 1.533inT 0.885
Length vs Age
L = 18,907 4 1.2tIT 0.941
TL = - 4,067 + 16,330InT OBIL
lunL = 2,789 + 0.0397 0.923
Inl, = 2,356 + 0.462InT 0,893
Weight vs Lengtt
W = | 221.04) ~ 7.499. 0.855
W ~ —614,020 + 320.40Mn1, 0.770
InW — (286 + 0,09XI 01.939
faW = — 6.7) + F.08SInL 0.966
a
LENGTH (m)
i)
On
.
ry *
z i] yrl5.907+1,211%
N=206
.
re 0.97 (P<0.001)
0 2 24 36 48
AGE (mo)
Fig. 3. Growth by length (cm) of captive A. antarcticus
over a four year period, Each symbol may represent
more than one data point,
Newly mature females measure approximately 30 cm in
length and males 44 cm’. In captivity these lengths may
be reached in ien and fifteen months, respectively, Hay
& Magnusson's death adders matured at similar ages to
wild populations’, Maturity is generally related to size
rather than age in repriles'¥:20_ This would explain why
some specimens held at Whyalla have been observed
mating at 19 mo}.
'Mirtschin, P. J. (1976) Pferpetofauna 8, 16-17.
2Hudson, P. (1979) Herpetofauna 11, 1113,
3Mirtschin, P. J. (1982) Herpetofauna 13, 14-17.
4 (1985) Jn G, Grigg, R. Shine & H, Ehmann
(Eds) “Biology of Australasian frogs and reptiles”. (Royal
Zoological Society of NSW, Sydney). pp. 505-509,
i) (1983) 8. Aust, Nat, 58, 24-28,
“Fitch, H. S. (1960) Herpetologica 16, 49-51,
‘Sokal, R. R. & Rohlf, FJ. (1981) “Biometry, 2nd
Edn”. (W. H. Freernan & Co., San Francisco),
§Fisher, R. A. (1930) “The genetical theory of natural
selection”, (Clarendon, Oxford).
°Saint-Girons, H, (1966) Mem. Inst, Butantan Simp.
Internac. 33, 105-114.
Hay, M. & Magnusson, W.
13-15.
'iCoyger, H. G. (1983) “Reptiles and Amphibians of
Australia, 3rd Edn”. (A. H. & A. W. Reed, Sydney).
Ek. (1986) Herp. Rev. 17,
G.R. JOHNSTON,
5005 F
2 1007
Ee Inye—6 7814 3, 08Sine
ion)
~~ N=206.
i 7=0,98 (P<D.009)
Ty
a) 7
gq 107
=
T 7 pt
0.05 0.1 0.5 1.0
LENGTH (m)
0,01
Fig, 4. Relationship between weight (gm) and length (cm)
in captive A. antarcticus. Each symbol may represent
more than one data point.
R. Angus, P. Fennell, P. Hudson, W, Ingall, P.
Mirtschin, H. Nygren, J. O’Reilly and H. Van Dyke helped
maintain the snakes upon which this study is based at the
Whyalla Fauna Park. K, Bradley, B. Doherty, T. Gruster
and M, Klessens assisted in taking measurements. Dr J.
Miller assisted with the analysis of growrh. Dr R. Shine
and Mr B. Roberts read dratt manuscripts.
1 Shine, R. (1980) Herpetologica 36, 281-289.
& Bull, it J. (1979) Am. Natur. 114, 296-303.
iptesmpiiene A. (1985) In Gi. Grigg, R, Shine & H.
Ehmanno (Eds). “Biology of Australasian frogs and
reptiles”, (Royal Zoological Society of NSW, Sydney).
pp. 185-192.
'SBull, J. J. (1980) Quart. Rey. Biol. 55, 3-21.
\6Heatwole, H. & Davison, FE. (1976) Herpetologiva 32,
332-336,
"Carpenter, C. ©, Murphy, LB. & Carpenter, G. C.
(1978) J. Herpetol. 12, 574-577.
Zar, J. H. (1974) Biostatistical Analysis (Prentice-Hall
Inc., Englewood Cliffs, New Jersey).
‘Porter, K. R. (1972) Herpetatagy (Saunders,
Philadelphia).
20Goin, C. 4. & Goin, O, B, (1971) Introduction to Herpe-
tology, 2nd Edn. (Freeman, San Francisco).
16 Nottingham Crescent, Glandore, South Australia $037,
PARATACAMITE FROM SOUTH AUSTRALIA
Summary
BRIEF COMMUNICATION
PARATACAMITE FROM SOUTH AUSTRALIA
Atacamite and paratacamite are important crystalline
modifications of copper(II) trihydroxychloride,
Cu,OH),;Cl, found in the upper portion of the oxidized
zones of many copper deposits throughout the world,
Atacamite, the orthorhombic polymorph of Cu,(OH),Cl,
has previously been reported from over twenty different
localities in South Australia, the most important being
the mines in the Moonta-Wallaroo district." In contrast,
paratacamite, the rhombohedral polymorph, has only been
reported from two localities in South Australia; Dome
Rock Copper Mine on Boolcoomata Station, Olary, and
the Mount Gunson Mine near Pernatty Lagoon. We now
report three new occurrences of paratacamite in this State;
see Fig. 1,
et med
Fig. 1. Map showing paratacamite localities in South
Australia.
Small, well formed, dark-green rhombohedral crystals
of paratacamite (Fig. 2) were found associated with
brochantite and quartz in cavilies ina brecciated hematitic
greywacke from the upper workings at Wallaces Gully gold
Prospect, 10 km NNW of Mt Painter in the Flinders
Ranges. A small crystal fragment was examined by single
crystal X-ray diffraction methods with the use of an Enraf
Nonius CAD4F diffractometer fitted with MoKa (graphite
monochromator) radiation. Accurate unit cell dimensions
were obtained from a least-squares refinement on the
setting angles of twenty-five well-centred reflections in the
range 7<#<22°% The cell obtained had parameters
a=9.169(6) A and @=96.25(7)" (hexagonal setting
a-13.659(6) and b~14.047(5) A), The mineral was
identified from its cell constants using the NBS Crystal
Data Identification File incorporated in the Chemical Data
Fig. 2. Paratacamite crystals on quartz from Wallaces
Gully gold prospect, 10 km NNW of ME Painter,
Flinders Ranges (Specimen SAM G13208).
Service VAX computer system, operated by the Science
and Engineering Researeh Council at Daresbury in the
U.K. The crystal structure of paratacamite has been
reported}
The identification of paratacamite from Wallaces Gully
prompted a systematic examination of the atacamite
specimens held in the collection of the South Australian
Museum, Paratacamite was identified, by powder X-y'ay
diffraction methods, on six specimens from the New
Cornwall Mine, Kadina, and on two specimens from the
Wallaroo Mines, also at Kadina.
At the New Cornwall Mine, paratacamilte occurs as line
grained masses honeycombed with vugs containing small
groups of atacamite crystals up to 1 mm in length, The
Coexistence of the two polymorphs indicates two periods
of crystallization, paratacamite being formed earlier than
atacamite. The formation of paratacamite is favoured by
relatively low concentrations of CuCl;.4 At higher
concentrations (greater than 0.04 F) atacamite is the
favoured crystallization product. Such variations in
conditions could be associated with movements in the level
of the watertable,
Paratacamite at Wallaroo Mines is associated with
aragonite, chalcocite and cuprite in an altered vein caleire
Here the paratacamite has been formed by the oxidation
of cuprite, The mineral occurs both as a fine crust
replacing cuprite crystals (Pig, 3) or as partially spherical
masses of small crystals, 0.5 mm thick, surrounding a core
of unaltered cuprite.
At the three localities cited, paratacamite appears to be
restricted to the upper part of the Oxidized zone,
We would like to thank Mr C, Johnston for providing
the material from Wallaces Gully and ta Ms F. Gommers
for assistance with powder X-ray diffraction and
128
photography. The financial support of the Australian
Research Grants Scheme is gratefully acknowledged.
'Noble, R. J., Just, J. & Johnson, J. E. (1983). “Catalogue
of South Australian Minerals.” (South Australian Dept.
of Mines and Energy, Adelaide.)
2Cloud, T. C. (1883). Trans. R. Soc. S. Aust. 6, 72-93.
3Fleet, M. F. (1975). Acta Crystallogr. B31, 183-187.
4Sharkey, J. B. & Lewin, S. Z. (1971). Amer. Mineral. 56,
179-192.
Fig. 3. Paratacamite replacing cuprite crystals with groups
of radiating aragonite crystals from Wallaroo Mines,
Kadina (Specimen SAM GI11409),
A, PRING, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000, M. R. SNOW and E. R. T. TIEKINK,
Department of Physical and Inorganic Chemistry, University of Adelaide, Adelaide, S. Aust, 5001.
PARASITES OF THE WOMBAT VOMBATUS URSINUS FROM GIPPLAND
REGION, VICTORIA
Summary
HRIEF COMMUNICATION
PARASITES OF THE WOMBAT VOMBATUS URSINUS
FROM THE GIPPSLAND REGION, VICTORIA
Lhere have been a number of taxonomic papers on the
helminths of the common wombat Forrhatus ursinus but
uo detailed studies of their parasite fauna. Fascioliasis,
an economically important disease of dumestic ruminants,
is sometimes found in wormbals grazing contaminated
aprigultural pasture.! But in o study on the prevalence of
Fasciola hepatica infection in south eastern Australia, na
hosts were collected from the Central Gippsland region.?
‘The womat is commonly and widely distributed across
Gippsland’, pasture damage and fence destruction being
such that it ts still legally designated vermin. From 1982
to 1984 wombat road kills were collected and examined
[0 determine their parasite fauna, and the results are
presented here.
Tiys, b. Localities in the fiippland tezion Where wombuts
were collected.
Although a number of road kills wece sighted within
(he study area, only seven were suitable for further
cyamination (Fig, 1). The internal viscera were examined
for gross pathological changes, then the lungs, liver and
gut contents were examined under a dissecting microscope
hot internal parasites. Nematodes. and castodes were tired
in 10% formalin, Nematodes were stored in 70% aleohol
with 5% glycerol and identified in Jactophenol, Ticks were
stored in 70o. alcohol.
Of the seven individuals examined, one was infected
with the mite, Sarcoptes scabiei Degeer, 1778 while five
harboured licks, Aponomme auruzinans Schultz, 1936.
No gross pathological conditions were found and no
helminths were observed in sites other than the digestive
tract. ‘lwo hosts harboured cestodes, Phascolotenta
comait Beveridge. 1976 in the ileum of one and
Triplotwenia undosa Beveridge, 1976 1n the duodenum of
another. All seven hosts were infected with the nematodes
Phascolostrongvlus turfevi Canavan, 1931 and
Vesophugosteumoides longisnicularis Beveridge, 197
while four hosts also were infected with Q. giltneri
Schwiitrtz, 1928 (Table 1). The parasite burdens were nol
high and except for the wombat with mange all the hasts
seemed to be in pood physical condition, No treniatode
infections, and more specitically no Fusciola hepatica
infections were found.
This is the first record of 7 undosa, previously known
only from macropodids, in a Wombat. Tripluteenia undosa
is found in a number of macropodid hosts. including
Wallabia bicolor and Macropus,fuliginosus, both of these
being cormmon throughout Victoria. However, the only
record of 7. wndasa from Cippsland hosts is trom We
hicolor collected ar Bonang in East Gippsland, and not
from either We bicolor or M. fitliginasus sharing their
range with wombats in the present study area
Ovsophagostomoldes silineri was the least common. of
the nematode species found and had not previously been
noted in hosts from Central Gippsland, Usually recorded
as parasites of the colon all three species were also found
in the small intestine and on one occasion rhe caccum,
This distribution is probably a function of post mortem
migration from) the colon. Although there is moderare
resistance by wombats to F heparica!, it is more likely
io this-case that no infections of F hepatica were found
because the specimens collected were not grazing in
infected areas. Central Gippsland is not one of the main
habitats for Lyymnaena tementosa the intermediate host
snail,> and nor is fascicliasis a siznificant problem of
sheep and cattle from the study area (Regional Vererinary
Laboratory, Bairnsdale, pers. comm.)
My thanks to 1. Beveridec for identifying the cesrodes
aud D. Kemp for identifying the ticks.
TABLE |. Sité in host anid abundance af infections with Oesophagostomvides longispicularis, ©: gilrneri aa
Phascolostrongylus turleyi from seven worsbars.
Site in Host
ileum Caecum Coton
Wo. Parasites/Host
Parasite Mean Range Mean Range Mean Rage
OL longispicularis 7 0.58 0.71 n-§ 23.7 0 62
O17 giltaert 54 0-29 6.14 a1 3.7 0-15
F turleyi 4a 0-294 0 a 47.7 0-113
Lee ccc nee te: ae
130
'Doube, L. J. (1982) Proc 4th Int. Conf. Wildl. Dis. Beveridge, I. (1976) Aust. J. Zoot. Suppl. Ser. No. 44.
Assn. Sydney 1981, 63-75. 5Seddon, H. R. (1967) “Diseases of Domestic Animals
2Spratt, D. M. & Presidente, P. J. A. (1981) Aust. J. Exp. in Australia. Part | Helminth infestations.” Revised by
Biol. Med. Sci. 59, 713-721. Albiston, H. E. (Commonwealth Department of Health,
3Norris, K. C., Mansergh, I. M., Ahern, L. D., Belcher, Canberra).
CA. A,, Temby, I. D., Walsh, N. G. (1983) Fish. Wildl.
Div. Vic. Occ. Paper 1.
LESLEY SMALES, Biology Department, Capricornia Institute, Rockhampton, Qld 4702.
THE OCCURRENCE OF DESMONEMA GAUDICHAUDI (LESSON)
(SCYPHOZOA, SEMAEOSTOMEAE) IN SOUTH AUSTRALIAN WATERS
WITH RECORDS OF FISH-JELLYFISH SYMBIOSES
Summary
GRIEF COMMUNICATION
THE OCCURRENCE OF DESMONEMA GAUDICHALU/DI (LESSON) (SCYPHOZOA,
SEMAEOSTOMEAE) IN SOUTH AUSTRALIAN WATERS WITH RECORDS OF
FISHJELLYFISH SYMBIOSES
The genus Desynoneme L. Agassiz, 1662 (Scyphowve,
Semiaeastamede) currently contains three species.t I has
an Amrarcti¢ re sub-Antarctic distribution?) but is
unfecotded in Australlan waters (the record of 2 rosea
Apassiz & Mayer trom Largs Bay, & Aust, refers 10 @
species of (yanea — see comment at end of this note).
Iwo specimens of #2 gawidichandi (Lesson) are recorded
lwte for South Australian waters, Both were found in
symbiotic relationship with lishes. Other jellyfish-fish
symbioses frona southern and eastern Australian waters
are documented. Maserial is deposited in the South
Australian Museum (SAM). Numbers prefixed by A refer
ta registrations in the natebooks of RYV.S.
Desnonewia guudicN@uds (Lesson) (SAN H348 (A1156)).
Bell width to turnover edge |] cms same measurement
with bell taid Mad 14 cm; 10-[5 tentacles/graup. Coll.
S.A. Shepherd, 29-iv.1969, 1 m depth, West Island (N.
side), Encoumer Bay, With SAM F5615 {A1157), a juvenile
trevally Pyendocarenx dentex (Bloch & Schneider) tearlier
Usacarantx georgianus (Cuvier & Valenciennes)) “living
under the mantle”
D. gaudichaudi SAM A349
Bell width to turnover edge 13 cm; same measurement
with bell laid fat 15 cm; & 20 tentacles/group. Coll. W.
Pots and 1. Phillips, 19.31.1986, opposite Point Camphell,
Coorong. SAM F4616, with ¥ small hardyheads, tamaly
Alherinidae (partially decomposed), among dentacles
(possibly Atherinosome miicrestama (Ginther), the only
member of the family currently recorded from the
Covrongs),
Previous records of D. ganelichaudiwith “juvenile fishes,
mostly of the genus Trachurus”, family Carangidae, have
been recorded from New Zealand.
Symbioses between jellyfishes and fishes have been
reviewed comprehensively,” Further records from
Australian waters are as fellows:
{. Unnamed species in association with juvenile yellowtail
scad, Trachurus mecullocki Nichols.4?
2, Pseudorhiza haeckeli Haacke (Rhizostomiesé)
(a) ALOIS, coll. S. A. Shepherd, 141.1968, near surface,
Aldinga Reef, S, Aust, off “Aldinga drop-olt”), SAM
F5614 (A1OL7), 3. small Pseudocaranx dentex and several
Sirene menoni Kramp (Hydrozoa, Leptometusae} (A116)
im association with the rhizastame.
(b) SAM A350 (A1391). Coll.'S. A. Shepherd, 254.1973,
depth 3 om. West Island, 15-16 small PR dentex (four
'Larson, R. (1986) Biology of the Antarctic Seas XVI,
Anlarctic Res. Ser. 41 (3), 59-165.
“Kramp, P-L. (1961) J. Mar. Dial Assoc. LK. 40, 1-469.
sO Sullivan, 1 (482) ANARE Research Notes 4, 1-33,
‘Agassiz, A, & Mayer, A, G, (1898) Bull, Mus, Comp,
Zool. Harvard 32f2), 15-19.
*Ghover C.J. ML, unpublished data.
‘Kingsford, M., in Larson, R, (1986) Hiolopy of the
Antarctic Seas XVL. Antarctic Res Ser, 41(9),
Frontispiece and pp. 102,. 148.
IMansueti, KR. (063) Copeia (1963, 1), 40-8o.
sampled, SAM F3905), and One small amphipod in
asyouation with the jellyfish,
tc) Jellyfish identified but not retained, 230.1975, Glenelg,
S. Aust, with juvenile ®& dentex (SAM F4238).
3, Cetostylus moaseicws (Quay & Gaimard)
(Rhizostomeac) SAM H351 (A884), Bell width 27 cm,
Coll. R. V. Southeort, 7.1965, Pore Hacking, NSW,
with juvetiles of T7 meeccteliochi (SAM P5617 (A886). This
association has been figured previously, Plate 15!.
4. Chrpsaura sp. (Senvaeosiomeae). Coll. R. ¥. Southeort,
17,4.1980, “Cape Jervis Station” of S. Aust. Dept. Fisheries,
Gulf St Vincent, near surface, from F. KR. V. Joseph Verco,
(a) SAM H352 (A2305) with P dentex, SAM F561]
(A2315).
(b) SAM 4353 [A2306) with F dentex, SAM P5612
(A2316).
(co) SAM FI354d (42307) with one juvenile mosaic
lcatherjacket, Lwhalichthys mosaics (Ramsay & Ogilby),
SAM F5613 (423345), (illustrated previously, Plate 14!2),
{d) A231, with P deniex, SAM F5610 (A2317),
(There were 0-4 # dentex symbiotic with each
Chrysaore.)
S. Cranec copitiata (L.) (Semaeastomeae), with juvenile
leatherjackers (family Monaesnthidac) (lidstrared
previously, Plates 14, 16!).
6. Cofostyhus mosaicus. Bass Strait, with juveniles of E.
SHasateus
Namenclatorial note
Desmonema rosea Agassiz & Mayer, 1B9B,* fram Lares
Bay, South Australia, is clearly a Cyaneo and nol a
Dexsmonema, since it has both radial and concentric
musele strands in the subumbrella, andthe tentacle origin
sites are U-shaped, not linear. Mayec!* synanymized this
species with Cyanea annaskala von |endenfeld, including
in the synonymy also C rvwellerianthe Haacke, 1887,"
from Gulf St Vincent, South Australia, Kramp (1995)!4
concluded that both of these last-named species are
synonyins of C expillva(l.). Womay be safely stated that
in Gull St Vincent there occurs one species of Cyanea,
C. capillata (L.). By transferring Lt rosea to Cyane in
1910, Mayer created a potential secondary hontonym of
Crenea rosea Quoy & Guimard, 1824, from the Grear
Harrier Reef, which Kramp*4 has regatded ns a doubtful
species al Craned, but possibly also a synonym of C
capitate. Thus four synonymy are available for C: rosea
{Agassiz, & Mayer).
*Seott, T. DE (1962) “The marine and fresh water fishes
of South Austria.” Handb. Flora & Fauna S. Aas.
(Gove Printer, Adelaide}.
Scott, T, D., Glover, C J, M. & Southoott, RY, (t974}
“The marine and freshwater fishes of South Australia"
(Second Edition). tfandb. Flora & Fauna 8. Aust. (Govt
Printer, Adelaide).
"Southear, ® ¥. (1982) Jellylishes (Classes Scyphozoa
aul ifyikrozoa). pp. 15-59. fn Shepherd, S. A. &
Thamias, T. M. (Eds) *Marine invertebrates of southern
132
Australia.” Part I. Handb. Flora & Fauna S. Aust. (Govt 12Mayer, A. G. (1910) Medusae of the World. Publicn.
Printer, Adelaide). No. 109, Carnegie Instn.
‘Last, P. R., Scott, E. O. G. & Talbot, F. H. (1983) !3Haacke, W. (1897) Jena Z. Naturw. 20, 588-638.
“Fishes of Tasmania” (Tasmanian Fisheries Development !4Kramp, P. L. (1965) Trans. R. Soc. S. Aust. 89, 257-278.
Authority, Hobart).
R. V. SOUTHCOTT, 2 Taylors Road, Mitcham, S. Aust. 5062 and C. J. M. GLOVER, South Australian Museum,
North Terrace, Adelaide, S. Aust. 5000.
VOL. 111, PARTS 3 & 4
30 NOVEMBER, 1987
Transactions of the
Royal Society of South
Australia
Incorporated
Contents.
Barker, S. Eighteen new species of Stigmodera (Castiarina) (Coleoptera: Buprestidae) - 133
Beveridge, I. & Sakanari, J. A. Lacistorhynchus dollfusi sp. nov. (Cestoda:
Trypanorhyncha) in elasmobranch fishes from Australian and North
American coastal waters - - - - - - - - - 147
Christophel, D. C. & Greenwood, D. R. A megafossil flora from the Eocene of Golden
Grove, South Australia - - - - - - - - - 155
Beveridge, I. & Campbell, R. A. Trimacracanthus gen. nov. (Cestoda: Trypanorhyncha:
Eutetrarhynchidae), with redescriptions of T: aetobatidis (Robinson,
1959) comb. nov. and 7, binuncus (Linton, 1909) comb. nov. - - 163
Geddes, M. C. Changes in salinity and in the distribution of macrophytes, macrobenthos
and fish in the Coorong Lagoons, South Australia, followig a period
of River Murray flow - - - oe - - - 173
Dulhunty, J. A. Salina bed instability and geodetic studies at Lake Eyre, South Australia - 183
Campbell, R. A. & Beveridge, I. Floriceps minacanthus SP. nov. (Cestoda: ba cat
from Australian fishes - - - - - - 189
Campbell, R. A. & Beveridge, I. Hornelliella macropora (Shipley & Hornell, 1906) comb.
nov. (Cestoda: Trypanorhyncha) from Australian elasmobranch fishes
and a re-assessment of the family Hornelliellidae - - - - 195
Brief Communications:
Davies, M., Watson, G. F. & Miller, C. A. New records of Uperoleia (Anura: Lepto-
dactylidae) from Western Australia with supplementary osteological
data on Uperoleia micromeles - - - - - - “ - 201
Reay, F. Australian plant nematodes: Longidorus Micoletzky, 1922 and Paralongidorus
Siddiqi, Hooper & Khan, 1963 (Nematoda: Dorylaimida) - - 203
Sokol, A. Yabbies at Dalhousie Springs, northern South Australia: morphological evidence
for long term isolation - - - - - - - - - 207
Alley, N. F. Middle Eocene age of the megafossil flora at Golden Grove, South Australia:
preliminary report, and comparison with the Maslin Bay flora - - 211
Errata:
Smales, L. Parasites of the Wombat Vombatus ursinus from The Gippsland Region,
Victoria - - - - - - - - - - - 213
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SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000
OF
TRANSACTIONS OF THE
ROYAL SOCIETY
SOUTH AUSTRALIA
INCORPORATED
VOL. Ill, PART 3
EIGHTEEN NEW SPECIES OF STIGMODERA (CASTIARINA)
(COLEOPTERA: BUPRESTIDAE)
BY S. BARKER*
Summary
Eighteen new species of Stigmodera (Castiarina) are described: S. aglaia sp. nov., S. alecgemmelli
sp. nov., S. arida sp. noy., S. atra sp. nov., S. caillaina sp. nov., S. chamelauci sp. nov.,
S. duggunensis sp. nov., S. earina sp. nov., S. gardnerae sp. nov., S. gordonburnsi sp. nov.,
S. goudiana sp. nov., S. jimturneri sp. nov., S. kanangara sp. nov., S. lauta sp. nov., S. livida sp.
nov., S. sundholmi sp. nov., S. vanderwoudeae sp. nov. S. viridissima sp. nov.
KEY WORDS: Coleoptera, Buprestidae, Stigmodera (Castiarina), New species.
FIGHTEEN NEW SPECIES OF STIGMODERA (CASTIARINA) (COLEOPTERA;
BUPRESTIDAE)
by 5S. BARKER*
Summary
Barker, S. (1987) Eighteen new species of Stigmodera (Custiarina) (Coleoptera: Buprestidae). Trans.
R. Soc. 8. Aust. 111(3), 133-146, 30 November. 1987,
Eighteen hew species of Stigmodera (Castiarina) are described: §. aglaia sp, nov., S. alecgemmelli
sp. noy., 5. arida sp. nov., S. atra sp. nov., S. cai/laing sp. nav., S. chamelauci sp. nov., 5. dugganensis
sp. nov, 5, earirta sp. Nov, S. gardnerae sp. noy., 8. gardanburnsi sp, nay, 3. goudiona sp. noy,, 8. jimlurneri
sp. nov., S. kKanangara sp. nov, §. lauta sp, nov., 8, livida sp. nov., 5. sundholihi sp, nov, 5, vanderwoudeae
sp. nov, 5, viridissima sp. nov.
Key Worps- Coleoptera, Buprestidae, Siégmodera (Castiarina), New species,
Introduction
Fighteen new species of Stigmodera (Castiarina)
are described herein and placed in species groups.
This brings the number of known species to about
four hundred. Most of the new species have
emerged from recent collecting, however several
have been known for a long time, but confused with
other species. All of them have been placed in
species groups on the basis of the structure of their
male genitalia and external morphology. All
previously recognised species groups are listed in
Barker (1986); an additional three groups are
detailed here.
Materials and Methods
Male genitalia were dissected from selected
specimens, cleaned of attached muscle and glued
dry onto a piece of card before they were photo-
graphed and printed at the same magnification.
Genitalia of known species are included for
comparative purposes, The abbreviations used in
the text for museum and private collections are as
follows (Watt 1979): ASSA Mr A. Sundholm,
Sydney, N.SW.; BMNFE! British Museum (Natural
History), London; EAQA Mr E, E,. Adams,
Edungalba, Qld; GBYA Mr G. G, Burns, Morning-
ton, Vic; JGAA Dr J. Gardner, Adelaide, 8. Aust.;
JHAQ Mrs J. Harslett, Amiens, Qld; JSBQ Mr J.
Sedlacek, Brisbane, Qld; ITNA Mr J. R. Turner,
Hazelbrook, N.SW.; MHSA Mr T, M, 38. Hanlon,
Ryde, N\SW.; MPWA Mr M. Powell, Attadale,
W.A.; NMVA National Museum of Victoria;
MMSA Macleay Museum, Sydney; OMBA Queens-
land Museum, Brisbane; RMBB L’Institut Royal de
Sciences Naturelles de Belgique, Brussels; SAMA
* Department of Zoology, University of Adelaide, G.P.O,
Box 498, Adelaide, S. Aust. SDOT.
South Australias Museum, Adelaide; WAMA
Western Australian Museum, Perth.
Stizmodera (Castiarina) alecgemmelll sp. nov,
FIGS 1A, 2D
Holotype; o, Stanthorpe, Qld, Deo. 1958,. +4.
Gemmell, SAMA 1 21178.
Allotype; 9, Stanthorpe, Qld, E. Sutton, SAMA
I 21179,
Paratypes: Qld: 1 o', same data as holotype; | co,
1 9, Stanthorpe, 27,x1.1926, S. M4. Haison, SAMA;
1 9, Stanthorpe, Nov. 1925, SAMA; | cr,
Stanthorpe, E. Sutton, SAMA; 1 9, Glen Aplin,
26.xi.1941, A. Gemmell, FHAQ; | o, 2 99,
Amiens, 26.xi1.1984, .( Harsleitt, JHAQ; t oc,
Fletcher, E. Sutton, SAMA; 1 @, Stanthorpe, &
Sution, JSBQ; 3 9 9, Stanthorpe, 202t1 1981,
Turner, JTNA; 1 of, Milmerran Dec, 1947/Jan.
1948, Macqueen, ASSA. N.SMW,: 1 o&, Coona-
barabran, 7.xi,1975, S. Barker, SAMA; 1 9,4km
E Rocky Glen, 3.%i,1981, 8, Barker, P Kempster, H-
Vanderwoude, SAMA; i o, 6 km SW Rocky Glen,
3,xi,1981, S. Barker P G. Kempster, H.,
Vanderwoude, SAMA; 2 ao @, 9 km N Coona-
barabran, 5/6.xi1.1983, A. Sundhalm, ASSA; 1c,
8 km N Coonabarabran, 8.43.1983, 4. Sundholm,
ASSA; 1 o, 7 km N Coonabarabran, 5.xi.1982, A.
Sundholm, ASSA; | co, Coonabarabran, 6.x1,1982,
A. Sundholm, ASSA; 2 oo, 1 9, 5 km WN
Coonabarabran, 4.x7.1983, 4. Sundholm, J, Bugeja,
ASSA,
Colour: Head, antennae, pronotum dull green.
Scutellurn blue. Elytra yellow with following black
markings with blue reflections: narrow basal
margin; broad pre-medial fascia not reaching
margin; broad posi-medial fascia reaching margin,
projecting anteriorly in centre of each side of
atiterior margin; pre-apical mark covering apex, all
S. BARKER
134
NEW SPECIES OF STIGMODERA 135
tnarks conilient along suture. Ventral surface and
tees dull green Hairs silver
Shape and sevlprvre: Head closely punctured,
inedian sulous, shart muzzle Antennae, seements:
1-3, obconic; 4, half toothed; 5-11, toothed,
Pronotum closely punctured, very small basal fovea
extending to apical margin as glabrous line, basal
natches on each side more lateral than medial;
apical margin projecting medially, basal margin
almost straight; laterally parallel-sided at base,
angled inwards, rounded to widest part before
centre, rounded and narrowed to apex, Scutellum
sculiform, glabrous, excavate, Each elytron
punctate-striate, intervals convea, heavily punctured
laterally and apically, smooth medially; laterally
angled out from base, rounded at humeral callus,
concaye, rounded posteriorly then narrowed io
bispinose apex; both spines small, margin rounded
between, apices diverging. Ventral surface with
shallow punctures, edges of abdominal segments
glabrous, dense medium length hairs, 87: truncate
in males; rounded In females,
Size Males, 13,040.24 » §5,3+6,09 mtn (17).
Females, 14.240.28 » 5.84013 mm (J)
Male genitalia; (Fig, 1A), Parameres more or less
parallel-sided for most of length, slightly widened,
rounded apically. Median lobe blunt, sides angled
away, Apophysis of basal picce medium width,
angled to apex.
Remarks: Member of S. simufata C & G species
group. Has been confused with 8. Durchetli C & G
(Holotype, MNHN). Aedeagus of S, burchellii C &
G (Fig. 1B) similar shape but longer and narrower.
Apex of median lobe puinted, sides acutely angled
away, Apophysis.of basal picce medium widrh but
not tapered apically. S. durchelliis coastal species
with ted margins on elytra. S, elegentnelli is found
inland and does not have red margins. Named after
the late Mr A. Gemmell, Glen Aplin, Qld,
Stigmodera (Castiarina) arida sp. 15v.
FIGS IC, 2B
Hotlolype co, Vic. NMYVA,
Allotype: 9, Sofala, N.SW., 26-x1.0083, A.
Sundaolm, SAMA | 21180.
Paratypes: Vie; 1 oo, 3 9 9, NMVA.
Colaur> Head dark blue, Antennae, segments: |, 2,
dark Blues 3-11, bronze, Promotum dark blue
medially, red laterally. Scutellum dark blue, Elytra
red with following dark blur clytral markings:
narrow basal margin; very broad pre-medial fascia
nov reaching margin, but cxpanded anteriorly to
basal margin, enclosing basal spot on each side:
very broad post-medial fascia expanded anteriorly
from centre of anterior margin on cach side and
on suture, expanded posteriorly from ventre of
posterior mangin on cach side and on suture; mark
covering apex. Ventral surface: pre-sternum variably
dark blue medially, red laterally; meso-and meta-
sternum dark blue or variably dark blue and red
laterally; abdomen red, Legs: femora and ubiae
bluc; tarsi blue-green. Hairs silver,
JShupe and sculpture: Head closely punctured, broad
median sulcus, short muzzle. Antennae. sepinenta:
1-3, obconic; 4-11, toothed. Pronotum closely
punctured, basal fovea extending to apical margin
as impressed line: apical margin projecting medially,
basal murgin barely bisinuate; laterally angled
oulwards from base, rounded to widest part
anteriorly, rounded and nartowed to apex, dorso-
ventrally fattened for one third of distance from
base. Scutellum scutiform, glabrous, extuvate. Each
elylron ptinctale-striale, intervals convex, punctured
more heavily laterally than medially; laterally
slightly angled our from base, rounded at humeral
callus, concave, rounded posteriorly then tapered,
rounded to spincless apex; minute indentation frorn
apical Margin lo suture without obyious spines,
apices diverging slightly, apical margin rough.
Ventral surface with shallow punctures, edges of
abdominal segments glabrous, sparse very short
hair. Sy: narrowly truncate in males; rounded and
pointed in females.
Size: Mates, 12,7 « 4.9 mm (2). Females 13.4
3.1 mm (4).
Male genitalia; (Fiz, 1C). Parameres parallel-sided
hasally, rounded posteromecially, slightly widened,
rounded apically. Median lobe sharp, sides acutely
angled away, Apophysis of basal piece short atid
slender.
Remarks: A member of S. distinevenda Saunders
species group. External morphology mast closely
resembles §. pilitaris Carter (Holotype male,
RMBEB), bui male genitalia are different. In S.
Fig. 1. Photomicrographs of male genitalia of (he following Srigmeodera (Castiarines species: A. §, aleceemmelli sp.
nay, B.S burcheili C & G, C. 5, avide sp, nov, D, 4, miliaris Carter, E. S. dugganensis sp_ nov., &. $. cehamelauel
sp. nov., G. S, vanderwovdere sp. nov, H, & gordumburnsi sp. nov. LS. klugiC & G, J, S, caillaina sp. nev.,
K. & goeudiona sp. nav, L. &. ‘vida sp. nov., M. §. wridissima sp. now., N, & earine sp. pov,, O, S. sundhofmit
<p. nov., P. S. imitetor Carter, Q. S. agiaia sp. now, R.S, gewtilis Kerremaus, 5. Satta ap now, To oblique
rremans, LS. geeeilior Carter, V. S gardnerae sp. ner, W, S reustelarsajar Thomaon, X, & enesma Cartes,
¥. S, junimeieri sp. aov., Z, keéenyvdre sp, niv,
RTH & BARKER
militariy (Fig. UD) paranieres are wider at apex,
median lobe blunt und obtusely angled away. Basal
piece is pot circular bul protrudes in centre,
Apopliysis of basal piece narrow, constricted in
middle and elongate. §. miditeris is stnallet species
with blue and yellow markings. The name refers to
the habitat and is derived from aridus L., dry.
Stigmodera (Castiarina) dugganensis sp. novi
FIGS IE, 20
Holotype: a, Duggan, W.A,, 27.46.1974, Kok i
Curhaby, SAMA | 21178,
Affotype; &, Lake Cronin, W.AL 17.x.198], M4.
Powell, WAMA.
Paratype WAs 2 oot, 80km B Hyden, 28.x.1984,
Af. Powell, MPWA! 1 cr, Lake Cronin, 18.x.J98!,
Af. Powell, MPWA, Vic; 1 ct, 1 @, Sea Lake, Nov.
Wh, Ho Gaudie, NMVA.
Colour Head blue. Antennae bronze-green.
Proneotum ble medially, red Eaterally from one
third of clistance from base ro apex, hase hve,
Scutellum bluc. Elytra red with (nllowing dark blue
markings: narrow basal margin; pre-medial fascia
not reaching margin but eatending anteriorly to
basal margin, enclosing hasal spot on each side;
broad post-medial fascia; mark covering apex,
marks mity or may not be conMuent along suture.
Ventral surface: sternum dark blue; abdomen blue
in centre, red at-sides, Legs: femora and tibiae blue-
freen; tarsi green, Hairs silver,
Shupe and seulplurer Head closely punctured, broad
median sulcus, short muzzle, Antennae, segments:
1-3, obcomie; 4-11, toothed, Pronotum closely
pinctured, narrow basal fovea extending to anterior
margin as thin impressed line, small basal notches
on each side more lateral than mediak apical margin
projecting medially, basal margin bisinuiles laterally
rounded from base (o widest part posreromedially,
Tamed to apex, Scutellum seutifarm, glabrous,
flat. Each elytron punctale-striate, intervals convex,
purtetured and wrinkled; laterally parallel-sided at
base then angled out, rounded at humeral callus,
faintly concave, rounded posleriarly.and narrowed
to Bispmose apes; small blunt marginal spine,
minule sutoral spine, anargin iidenied between,
apices diverging slightly, Ventral surface with
shallow punctures, sparse medium length hairs: S32
truncate in males; rounded and slightly pointed in
fernales,
Size: Males, 12.3 = 4,5 mm (5). Vemales, 12.9 +
SAY ay (2).
Mate zenitaliq: (Fig, VE), Paraincres parallel-sided
hisally, widened postérdmedially, mare or less
parallel-sided, rounded apically Median. lobe
pointed, sides acutely angled away. Apophysis of
basal piece narrow and tapered apivally.
Remarks: Mecober of the S. distinguenda Saunders
species group on basis of pronotal markings,
external morphology and male genitalia, Although
distinct within its specics group, could be confused
with & rufa Barker, a member of the §. pibbicollis
Sanders species group, which has sintilar colour
and pattern. Male genitalia are very different Barker
(1986, Fig, 2E), The name is derived from the type
toculity.
Sligmodera {Castiarina) chamelauci $p. nov,
FIGS IF, 2P
Holotype: o, Spalding Park, Bluff Pi, Geraldton
District, W.A., $.ix.1973, AN. McForland, BMNH.
Altaiype: 9. Spalding Park, Bluff Pr, Geraldton
District, W.A_, 21.14.1973, NV. McFarland, BMNH.
Peratypess WAL 25° oc, Dongarra ftsic),
6-19.5x.1934, 20-25.ix.1935, 26 ix-3,.x.1935, 4-10.x,
£935, 11-28.%,1035, R, E. Tuer, BMNH & SAMA,
Cotour; Head dark bluc. Antennaé, segments: 1,
blue; 2-11, blue-green, Pronotum dark blue with
bronze reflections, Scutellum dark blue. Elytra
orange with rcd margin and following black
markings wilh blue reflections: narrow basal
margin; pre-medial fascia not reaching margin or
represcated by spot over each humeral callus and
one on suture; post-medial fascia reaching, margin,
expanded anteriorly on suture and on anterior
margin on each side more lateral than medial,
expanded posteriorly on suture and on posterior
margin oneach side more lateral than medial; pre-
apical mark also covering spines, confluent ip more
heavily marked specimens, post-medial fascia and
apical mark con!Iwent along suture. Ventral surface
dark blue. Legs: lenvora and tibiae dark blue; tarsi
blue-green, Hairs silver.
Shape and seulpture: Head Sosely punctured, broad
median sulcus, short muzzle. Antennac, segments:
1-4, ubgenic; 3-11, toothed. Pronotum closely
punctured, narrow basal fovea extending to apical
margut as glabroys line, basal notches represented
by glabrous area on cach side more lateral than
medial? apical margin straight, basal margin
bisinuate; laterally rouoded from base, widest
postcromedially, rounded and narrowed tu apex,
Scutellum scutiform, glabrous, excavate, Each
elytron punctate-striate, intervals convex, punctured,
More $0 lalerally than medially; laterally angled out
frum base, rounded at humeral callus, concave,
rounded posteriorly and narrowed, rounded again
to bispittose apex; sharp marginal spine, sniall
sutiiral sping, margin rounded and mdented
NEW SPECIES OF STIGMODERA WwW
Fig. 2. A, Stigmodera gordonburnsi sp. nov., B.S. arida sp. nov., C. 8. duggunensis sp. nov., D. S. ulecgernmtelli
sp. noy., E. S, caillaina sp. novy., F. S. gardnerae sp. nov., G. S. livide sp. nov,, H- 8. viridissima sp. nav., 1. S. earinta
sp. nov, J. &. fauta sp. nov, K..S. a/ra sp. nov, L, S. sundholmi sp. noy., M.S. aglaia sp. nov., N.S. jimfurnert
ap. nov, O, S. vanderwoudeae sp, nov., P, S, chomelauct sp. nov. Q. §. goutliana sp. noy.. R. 5. kunangara sp, nov,
LR 5S. BARKER
betwoon, apices diverging, apical margin subserrate.
Ventral surface with shallow punctures, edges of
whdominal segments glabrous, moderately hairy,
hairs medium Jength. $7: truncate in males:
truncate ancl indentec! medially in females, Legs,
male: tarsal pads absent on Jegs 2 and 3 on
tarsomeres 1-3, replaced by median spine
Sizes Males, l0.8+0,)3 » 4.146.065 mm (26).
Female, 119 x 4.5 nm (i).
Maje genitalia; (Fig, IF) Wedee-shaped with
relatively narrow basal piece, apophysis and
purameres,
Remarks: Member of S. sexplagiata Gory species
group on basis of iis wedge-shaped male gentlalia
and external morphology. Elytral colour and red
margins distinguish this species from others in
group. Named afier Chamelaucium uncinatum
Schau., Geraldton wax, on which adulis are
collected,
Stiguindera (Castiaring) vanderwoudcav sp, nov.
FIGS IG, 20
Hotetype: 0, Lake Gilles Conservation Pk, 27 km
E Kimba, S. Aust,, 25.x.1982, S. Barker PG.
Kempster, H. Virtderwoude, SAMA 1 21182,
Atotype: 2, sane dataas holotype, SAMA 1 21183,
Paratypes: S. Aust 1§ oor, 8 9 9, same data as
hnlntype, SAMA.
Cofuur; Head black with blue reflections.
Antennae, scements:! 1, 2, blue; 3-11, blue-green.
Prunolum and scutellum black with blue
reflections, Elytra orange with following dark blue
markings: narrow basal margin; pre-medial Fascia
not reaching margin, projecting slightly anteriorly
on sutwe; broad post-medial lascia projecting
anteriorly on sulure at anterior matgin and in centre
af each side of anterior margin, projecting
posteriorly on suture at posterigr margins mark
covering apex, murks confluent along suture,
Ventral surface and legs dark blue. Hairs silver.
Shape und sculpture; Head closely pynetured,
median sulcus, short muzzle, Antennae, segments:
1-4, 1obconic; 5-11, toothed. ronotum closely
punclured, narrow basal toveu extending to centre
as glabrous line, then to apical margin a3 impressed
line, apical margiu straight, basal margin bisinuate;
laterally angled inwards from base, rounded to
widest part posteromedially, rounded and narrowed
to apex. Scutellum scutiform, glabrous, excavate,
Each ¢lytron punctate-striate, intervals convex,
punctured more heavily laterally than medially:
laterally angled oul trom base, rounded at humeral
callus, concave, rounded posteriorly aud narrowed
to bispinose apex; sharp marginal spine, uinule
sutural spine, margin rounded and indented
between, apices diverging slightly, apical margin
subserrate, Ventral surface with shallow punctures,
edges of abdominal segments glabrous, moderately
hairy, hairs short, ‘S7 truncate in both sexes. Legs,
male: tarsal pads absent on fegs 2 and 3 on
tarsomeres I-3 each replaced by small double,
median spine.
Sizes Males, 7.940.270 = 3.0+0.09 mm [l6p.
Females, 8.64024 x 3.4+ 0.04 mm (9).
Male genitalia: (tig. 1G). Wedge-shaped. Parameres
parallel-sided at base, widened posteruniedially,
slightly rounded then abruptly rounded near apex,
apical edge concave. Median lobe pointed, rounded
apically, Apophysis of basal piece narrowed,
Remarks: Member of S. sexplagiata species grougr
on basis of its wedge-shaped male genitalia and
external morphology, Most closely resembles S$
curnishi Barker (1983, Fig, 1R) bur is smaller Male
gcnitalia are smaller and not as wide apically and
spines un ventral surface of tarsomeres are much
less prominent than in S. coraski, All specimens
were collected on flowers of Lremophila scoparia
(R.Br) F. Muell, Named after Ms H. Vanderwoude,
Adelaide.
Sigmodera (Castiariana) gordonhurnsi sp. nov-
TiGS 1H, 2A
Holotype: Oo, Grampians, Vic. 22.xi.1982, GL G.
Burts, NMVA.
Allotype @, Grampians, Vit, (9.xi.1983, G. GC.
Burns, NMVA,
Paratypes; Vic: } o, same data as holotype, GBVA;
1 &, Inglewood, Nov. 1952, & Smith, NMVA: 1 9,
same data ax dllorype, GRVA; | o, Grampians,
23.¥1.1982, GG. Burns, SAMA; 1 0, Grampians,
24.x1.1982, G. G. Burns, GBVA; 1 9, 16 km N
Benambea, 14./.1979, PL Cullen, SAMA,
Colour: Head dark blue, Antennac, segments! 1, 2,
blue-ereen; 3-11, bronze-grocn. Pronotum dark bie
medially, blue laterally. Scutellurn dark blue, Elytra
red with following dark blue markings: broad basal
margin, red spot at lateral margiir on cacle side;
broad post-medial fascia; mark covering apex, all
marks confluent along suture, Ventral surface blue.
Legs! femora blue; tibiae and tarsi bluc-greeti. Hairs
silver,
Shape and sculpture: Head closely punctured, broad
median sulcus, short muzzle, Antennae, segments:
1-3, obconic; I-(l, toothed. Pronotum closely
punctured, basal fovea extending (o apical margin
4 glabrous line, basal notches on cach side more
lateral than medial; apical margin projecting
medially, basal imargiit almust straight; laterally
NEW SPECIES OF STIGMODERA 139
parallel-sided at base, angled outwards, rounded to
widest part posteromedially, rounded and narrowed
to apex, Scutellum scutiform, glabrous, Mat. Each
elytron punctate-striate, intervals convex, punctured
and wrinkled; laterally angled out from base,
rounded at humeral callus, concave, rounded
postenorly to bispinose apex; both spines small and
blunt, margin rounded and indented between,
apices diverging slightly, apical margin subserrate.
Ventral surface with shallow punctures, edges of
abdominal segments plabrous, sparse short hairs,
S72 truncate In male; rounded in females.
Size: Males,16.2 « 6.5 mm (5). Females, 17,3 x
7.3mm (3).
Mfole gentiatia: (Fig. 1H). Parameres parallel-sided
basally, rounded posteromedially, widened, rounded
apically, Median lobe pointed, sides acutely angled
away then rounded and obtusely angled away.
Apophysis of basal piece medium width.
Remarks: Member of S. &iug? species group, In
closest species 8, kiugi C & G (Tig- 11), aedeagus
is shorter, apices of parameres are more abruptly
rounded, median lobe blunter, Only red and blue
Species in group, S. Afneiis yelow and blue and &
rubicunda Carter is ted and purple, Named after
Mr G. G, Burns, Mornington,
Stigmodera (Castiarina) caillaina sp. nov,
FIGS 11, 2E
Netotype: o, Vic, Blackburn, SAMA | 21184,
Allotype: >, Hornsby, N.SW., Schrader, SAMA |
21185.
Paratype: | or, no data, RMBB,
Colour: Head and antennae blue-green. Pronotum
green medially, blue laterally, Scutellum blue, Elytra
yellow with following black markings with blue
reflections: narrow basal margin; pre-medial fascia
teaching margin with projection from posterior
margip; posi-medial fascia reaching margin,
concave posteriorly; mark covering apex, all marks
contfuent along suture. Ventral surface dark blue.
Legs blue-green. Hairs silver,
Shape and sculprurer Head closely punctured,
median sulcus, shor muzzle. Antennae, segments:
1-3, obconic: 4-11 toothed, Pronotum closely
punctured, small basal fovea extending Lo centre as
glabrous line; apical margin slightly projecting
medially, basal margin barely bisinuate; laterally
angled outwards from base, rounded to widest part
postéromiedially, tapered to apex, dorso-ventrally
flattened on basal half, shallow foves at each basal
angle; Scutellum scutitorm, glabrous. Mat. Each
elytrim punctate-stnate, intervals convex, punclured;
laterally angled our from base, routrded at humeral
callus, concave, rounded posteriorly and narrowed
to bispinose apex: small marginal spine, small
sulural spine, margin rounded and indented
between, apices diverging, apical margin subserrate,
Ventral surface with shallow punctures, edges of
abdominal segments glabrous, moderately hairy,
hairs medium length. S7: truncate in both sexes,
Size: Miles, 14.9 » 6.2 mm (2). Female: 19.7
8.5 mm (1).
Male genitalia: (Fig, (J). Parameres parallel-sided
basally, widened posteromedially, parallel-sided
anteromedially, rounded apically, Median lobe
pointed, sides acutely rounded away. Apophysis of
basal piece narrowed,
Remarks: Belongs in 5, cupricollis Saunders species
group, Most resembles S. deyrollei Thomson but
male genitalia are smaller, apically wider and
apophysis more constricted chan in & deprolles
Barker (1986, Fig. 1M). 3. catllaine is smaller than
& deprollei, pronotum of S. deyrollel is green and
underside green not dark blue as in S, caillaina.
Name refers to colour of pronotun and is derived
from caiainus L., blue-green.
Stigmodera (Castiaring) goudiana sp. nov.
FIGS IK, 2Q
Holotype: x, Sea Lake, Vic,, Nov, 1916, HS Gotelie,
NMVA.
Allatype: 9, same data as holotype, SAMA [ 21184.
Paraiypes NISW.s L oc, Roto, 30.2.1981, JR,
Turner, ASSA; 2 oc, 1 km S Matakana,
30.x.1984, A, Sundholm, J Bugeja, ASSA, SAMA!
J 9, Roto-Matakana, 28.41.1981, A. Sundholm,
ASSA.
Colour: Head black with bronze and blue
reflections, muzzle blue. Antennae and pronotum
black with bronze and blue reflections. Scutellum
black, Elytra yellow with following black markings:
narrow basal margin; incomplete pre-medial fascia,
represented by mark on cach humeral callus and
one on suture confluent with basal margin around
scutellum; incomplete post-medial fascta repre-
sented by mark om cach side and one on suture,
confluent in some spevimens; niark covering apex
and spines, lasi two marks may or aay not be
confluent along suture, Ventral surface black with
bronze and blue reflections. Legs; femora and tibia
blue; tarsi bluc-green, Hairs silver.
Shepe and s¢cstipture; Head closely punctured,
median sulcus, short muzzle. Antennae, segitents:
1-3, obconic; 4-11, toothed. Pronatum shallowly
Punctured, narrow basal fovea, basal notches on
each side more lateral than medial; apical margin
straight, basal margin bisinuate; laterally rounded
Tay $. BARKER
from base, widest posteromedially, rounded and
narrowed to apex, Scutellum scutiform, glabrous,
excavate. Each clytron punctate-striaie, intervals
convex, punctured, lateral margin flattened; laterally
angled out from base, rounded at humeral callus,
suncaye, rounded posterlorly and narrowed to
bispinose apex; both spines sharp, margin rounded
hetween, apices diverging. Ventral surface with
shallow punctures, edges of abdominal segments
glabrous, sparse short hairs, 34: truncate in both
SEXES,
Sizer Males, 10.8 « 4.3 mm (4}, Females, 11.8 «
5.2 mm (2).
Mole genitelia: (Fig. 1K). Short. Parameres widened
basally, rounded posteromedially then widened,
rounded apically, Median lobe pointed, sides acutely
angled away. Apophysis of basal piece narrow.
Remarks: Member of S, delectahilis Hope species
group, closest to S. dessarti Barker (1986, Fig. 30)
from W_A. Paramcres in that species are narrower
und its has red elytra. Namved after late Mr W.
Goudie, Sea Lake, Vic.
Stigmodera (Castiarina) vida sp. nov:
FIGS IL, 2G
Holotype o, Bukey, Qld, Jan. 1934,F HW Hilsan,
NMVA,
Allotype: ?, MacPherson State Forest, 27 km NW
Central Mangrove, Qld, {7.xii.1979, £ & D. Gardner,
SAMA I 21187.
Paratypes Qld: 1 o, | >, Eukey, 29.1975, &
Barker, SAMA; 3 9 9, Glen Aplin, 28.x.1975, S.
Barker, SAMA; 2. 0'o, same data ay holotype,
NMVA; 1 0, 1 9, Mt Tambourine, 26.xi1.1937, G
Oke, NMVA3 1 9, Durack via Brisbane, 19,x.1978,
G. G. Burns, GRVYA; 1 9, Amiens, 9.xi17,1979,
T. M.S. Hanlon, MHSA; | 9, Stanthorpe, Dec.
1981, A. Sundholin, ASSA. N.SW.2.1 of, Grafton,
SAMA: 4 cu, 2.99, Armidule, Deg, 1953,
CG. L. Gooding, SAMA; 5 wo, 2 949,
Glenbrook, 7) J) Hawkeswood, SAMA: 5 oo,
2909, 96km SE Cessnock, 14,.xi1.1979, 1 & D,
Curdaer, IGAA: 14 oo, 6 9 9, Appin, Nov, C
Dveuquet, RMBB; 8 coor, 4 O 9, Glenreagh, Ocr.
1922, C. Deuguet, RMBB; I co, 4 99,
Mendaouran, Nov. 1933, C. Derequer, RMBB; 1 9,
73 km ENE Windsor, 18.xi,1984, 7 M.S. Hanlon,
MHSA;7 oor, 3 9 9, Dunnedao, L-xiL.1985, A.
Sundhoim, ASSA: 1 o, Sandy Pt, Sydney,
Lxii.1980, A, Sundhofm, ASSA; 1 co, Hassans
Walls, Lithgow, 28.xi.1982, JR. Turner, ASSA;
1 9, Hill End, 24.x1.1983, . . Tiermer, ITNA: I
9, Dangar Falls, 14,x37,1983, 0 R. Tiurrer, JTNA;
loo, 1 9, Fitzroy Falls, Jan. 1941, C2 Denguet,
MMSA, Vie: 2 oo, Rushworth, 7.91,1976, G, G.
Burns, GBVA; 1 cf, 1 @, Inglewood, 9,x1,1977,
G. G, Burns, GBVA,
Colour; Head and antennae blue. Pronotum
variably black medially, blue laterally. Scutellum
black. Elytra yellow with following black markings
with blie reflections: narrow basal margin; pre-
medial fascia not reaching suture but ¢xpanded
anteriorly over humeral callus and posteriorly
reaghing lateral margin in some specimens, not
expanded posteriorly in others; very broad post-
medial fascia; mark covering apex and spines, last
three marks confluent down suture. Ventral surface
and legs blue, Hairs silver.
Shape and sculpture: Head closely punctured,
shallow median sulcus, very shore muzzle.
Antennae, seements: 1-4, obeonic; 5-11, toothed.
Pronotum closely punctured, marrow basal fovea
extending to apical margin as glabrous line; apical
margin straight, basal margin bisinuate; laterally
parallel-sided at base, angled outwards and rounded
to widest part anteromedially, rounded and
narrowed apically, Scutellum scutiform, glabrous,
flac. Each elytron punctate-striate, intervals flat
medially, convex elsewhere, smooth; laterally angled
out from base, rounded at humeral callus, concave,
rounded posteriorly then Warrowed, rounded to
bispinose apex; small sharp marginal spine, very
small sutural spine, margin rounded and indented
between, apices diverging slightly. Ventral surface
with shallow punctures, edges of abdominal
segments glabrous, sparse short hair. 57: truncate
in males; rounded in females. Legs, male: tarsal
pads missing on all legs from tarsomeres | and 2,
each replaced by single median spine.
Size: Males, 7-5+0.08 x 2.8£0.02 mm (57),
Females, 4.14010 ~ 3.0+0,04 mm (36).
Male genitalia; (Fig. 1L). Parameres widened
basally, notched anteromedially, parallel-sided,
rounded apically, Median lobe pointed medially,
sides angled axway obtuscly. Basal piece medium
width.
Remarks: Member af 8. flavopicta (Boisduval)
species group, Most like §. Sicelar C & G from &
Aust., figured under its synonym S. aliceae Barker
(1980, Figs 20, 36), Apophysis of basal piece is
wider than in S. bicolor and median lobe sharper.
Also 8, bicolor has unicolorous pronotum, Often
misidentified in museum collections as S. flavavaria
Saunders (replacement name for & flavopicia C &
G, primary homonym of 5, flevapicta (Boisduval)).
Original description and illustration indicate that
S. flavevuria Saunders is known green morph of
S. flavopicia (Boisduval). Name refers to
hicalouraus prometuor and is derived from /iviedeus
L., black and blite
NEW SPECIES OF STIGMODERA 141
Stigmodcra (Castiarina) yiridissima sp. nov,
FIGS IM, 2H
Holotype; o, Dunmore State Forest, Old, 2.i.1985,
M, Powell, SAMA 1 21188.
Allotype: >, sane data as hulotype, SAMA | 21189,
Poralype; Qld; | o, same data as holotype, MHSA.
Colour: Head, antennae, pronolum and sculellum
green, Elytra yellow with following black markings:
narrow basal margin; pre-medial fascia not reaching
margin, expanded anteriorly to humeral callus;
post-medial fascia reaching marvin; mark covering
apex and spines, last three marks confluent along
suture, Ventral surface and legs green. Hairs silver.
Shape and sen(pture; Head closely punctured,
shallow median sulcus, very short nuzzle. Antennae
compressed, segments; t-4, obconic; 5-11, toothed.
Pronotum closely punctured, narrow basal fovea
extending to apical margin as impressed ling, basal
notches on each side more lateral than medial;
apical margin straight, basal margin bisinuates
laterally angled outwards trom base, rounded al
widest part anteromedially, rounded and narrowed
apically. Scutellum scutiform, glabrous, excavate.
Each elytron punctate-striate, intervals flat medially,
convex elsewhere, 3rd and Sth raised at apex, LOth
taised for most of its length: laterally angled oul
from base, rounded at humeral callus {widest part),
concave, rounded posteriorly and narrowed to
bispinose apex; small blupt marginal spine, minute
autural spine, margin indented between, apices
diverging, Ventral surface with shallow punctures,
edges of abdominal segments glabrous, moderately
hairy, hairs medium length. Sy: truncare in males;
truncate and tridented medially in females,
Size: Males, &8 x 3.4 mm (2). Female, 8.9 x
3.5 mm (1).
Male genitalia: (Fig. UM). Parameres parallel-sided
basally, rounded posteromedially, parallel-sided
anteromedialy, faintly notched, rounded apically,
Median lobe pointed, sides acuicly angled away.
Apophysis of basal piece wide,
Remarks: Member of S. eruentate (Kirby) species
group. Because of its colour, pattern, shape of
pronotum and raised intervals on elytra this species
is distinct within S. erwensere spectes group. Name
refers to colour ol head and body and is derived
from piridts L., green.
Stlgmodera (Castiarina) carina sp. nov.
FIGS IN, 2l
Holotype cr, 33 km N Israelite Bay, W,A.,
24.x.1980, S. Barker, P G, Kempster, WAMA,
Allotype- 9, same data us holotype, WAMA.
Paraiypes W.A2 9 oo, 3 99, same data as
holotype, WAMA & SAMA,
Coiour) Head, antennae, pronowum and scutellum
green ov bhie-green. Elytra Orange with Following
black markings with blue reRections: narrow basal
margin; broad pre-medial fascia, not reaching
inargin but expanded anteriorly to humeral callus
and posteriorly reaching lateral margin; broad post-
medial fascia reaching margin: apical mark, all
marks confluent along suture. Ventral surface and
legs green or blue-green, Hairs silver.
Shape and scuiplure: Head closely punctured,
shallow. median sulcus, shor! muzzle, Antennae,
segments: L-4, obconic; 5-11, toothed. Pronotum
closely punctured, narrow basal fovea extending to
near apical margin as impressed line; apical margin
straight, basal margin bisinuate; laterally parallel-
sided at base, rounded to widest medially, rounded
and narrowed to apex. Scutellum scutiform,
glabrous, excavate. Each elycron punctate-striate,
intervals convex, punctured; laterally angled out
from base, rounded at humeral callus, concave,
rounded posteriorly and narrowed to bixpinose
apex; both spines small, margin rounded and
indented between, apices diverging slightly. Ventral
surface with shallow punctures, edges of abdominal
segments glabrous, sparse short hair, Sj: truncate
in both sexes,
Size; Males, 9340.25 » 3.1+0,09 mm [1)),
Females, 9.74.0.28 » 3.310,08 mm (4).
Male genitalia: (Fig, 1N), Parameres parallel-sided
basally, rounded posteromedially and slightly
widened, faintly indented, rounded apically, Median
lobe pointed, sides acutely angled away. Apophysis
of basal piece median widih.
Remarks: Member of S. ervenrata (Kirby) species
group. Must resembles 5, moritigera Oke which bas
blunter median lobe and broader parameres (Barker
1983, fig, IW), S. otontigeria has bicolorous
pronotum and occurs in Australian Alps, Name
refers to colour of head and body and is derived
from eurines L.. green.
Stigmodera (Castiarina) sundholau sp. nov.
FIGS 10, 2L
Halaiype: of, 4m W Paluma, Old, 3.4,1986, BB,
& & WS Adams, SAMA 1] 21190.
Allatype: 9,4 km W Paluma, Qld, 77.1986, & E
&& Wes Adanis, SAMA 3 21191-
Peraivpes; Qld; 3 to, 2 90,4km W Palma,
41.1986, 4. Sundhoim, ASSA; 1 oo, 5/6/7U.
1986, Paluma, ££ & BOM Adams, POA
4c. 8 2, 4km W Paluma, 677/8/10.1,1986,
J Bugeia & A, Sundioim, ASSA.
42 S, BARKER
Colour: Head black wilh blue-green reflections,
Antennae, segments: 1, 2, blue-green; 3-11, bronze.
Pronotum black with blue-green reflections.
Seulelluin black woli Glue ceflevions. Elytra yellow
with following black markings; narrow basal
margin; pre-medial fascia reaching margin,
expanded anteriorly from anterior margin aver
humeral callus, enclosing elongate yellow basal
mark and variably an clongate yellow mark on
margin at humeral callus; post-medial fascia
reaching margin and expanded anteriorly From
antcnor margin, in some specimens fascia conflucnt
enclosing yellow mark in middle of elytra and one
on tateral margin; mark covenng apex and spines
with large mark between it and posi-medial fascia,
yellow vlosest to suture, red from centre Lo margin,
all marks broadly confluent along suture, Ventral
surface green with blue reflections. Legs: femora
and litsae blue; tatsi blue-green. Hairs silver.
Shape and sculpture: Head closely punctured,
median sulcus, short muzzle. Antennae, segmenrs:
1-3, obconic; 4 half touthed; 5-11, toothed, Pro-
notum closely punctured, basal fovea extending to
apical margin as impressed line; apical margin
straight, basal margin bisinuate; laterally widest at
base, rounded and narrowed to apex, elongate
shallow fovea at each angle, Scutellim cordiform,
punctured, excavate. Each elytron punctate-striate,
scurellary, 3rd at anterior part, Sth and 7th imervals
slightly raised and smooth, rest flat and heavily
punctured, lateral margin flattened; laterally angled
oul from base, rounded at humeral callus, concave,
rounded posteriorly and narrowed to bispinose
apex; small sharp marginal spine, minute sutural
spine, margin rounded and indented between, apices
diverging slightly. Ventral surface with shallow
punctures, edges of abdominal segments glabrous,
mpderately hairy, hairs short. Sj: truncate in
males; truncate and indented medially in fernales.
Size: Males, 9540.10 x 3.640,04 mm (28).
Females, 10,0 +0.20 x« 3.8+0,09 mr (8).
Male genitulla: (Fig 10), Parameres parallel-sided
basally, widened posteromedially, rounded apically,
Median tobe blunt, sides obiusely angled away,
Apopliysis ol! basal piece breve.
Remarks: Member of 5. Sella Saunders species
proup (listed under nexc species). Alittougty this
species resembles S. gentilis Kerremans, genitalia are
most like those of S. imttator Carter (Fig. 1P),
which, although broad upically are not spoon.
shiuped, Rlytral colour resembles 5, geasilis but
upper surtace of S. seneilis is heavily punctured,
while that of S. suadheda is glabrous und elvtral
patterns are quite differenr. Also & gensilis is much
smuller species than S, sundkolmi. Named after Mr
A. Sundholim, Sydney.
Silymuders (Custiaring) aglaia ep. nov.
FIGS 10, 2M
Holotype: oo, 4 ki W Paluma, Old, 4.1, 1986, A.
Suadhoim, SAMA J 2121).
Allotype: Q, same data as holotype, SAMA 1 21201,
Colour: Head black, muzzle blue. Antennae green.
Pronoturn and scutellum black with bronze reflec-
tions, Elytra yellow. with following black markings:
broad basal margin; broad post-medial. fasvia
angled anteriorly but reaching margin; apical mark,
all marks broadly confluent along suture enclosing
& pre-medial yellow mark on each side reaching
miarein, large red pre-apical mark along each
Margin, merging into yellow closest jo suture,
Ventral surlace green with geld reflections, Legs:
femora and tibjse blue; tarsi blue-green. Hairs silver.
Shape and sculoture: Head closely punctured,
median sulcus, short muzzle. Antennae, segments
l-4, obconic; 5-1), toothed. Pronotum closely
punctured, basal fovea extending to apical margin
as impressed line; apical margin projecting medially,
basal margin bisinuate; laterally parallel-sided at
base, rouided posteromedially and narrowed to
apex, round fovea on each side more hasal thac
medial, shallow fovea al each basal angle. Scutellum
seutiform, glabrous, excavate, Hach elytron
puncrate-stiiate, intervals convex, punctured lightly
from suture to 6th, punctured heavily from 7th
interval to margins laterally angled out [rom base,
rounded ai himeral callus, concave, rounded
posteriorly and narrowed to bispinose apex; sharp
marginal spine, small sutural spine, margin rounded
and indented between, apices diverging. Ventral
surface with shallow punctures, edges of abdominal
segments Blabrous, moderately hairy, hairs short.
$4: truneate in both sexes.
Sizes Male, 3,0 ¥ 8.5 mim (1).
Male genitalia: (Fig. (Q). Parameres widened
basally, rounded and narrowed apically, Median
lobe sharp, sides angled away, Apophysis of basal
piece moderately wide,
Remurks: Belongs in S, betla Saunders species proup
comprising & fella Saunders, 8. agiaia sp- nov, 4.
dedi Carter, S gentilis Ketremans, S. imitator
Carter, S. kerremansi Blackburn, §. kershawe
Carter, S. artrginata Barker, 5. sundholmi sp. now,
Closest to. gentilis Kerremans (Pig. 1R) which has
parameres more or less parallel-sided on apical half,
sides of qnedian lobe acutely angled away. 8. gerttilis
has four yellow spots on each elyiron while §, aeleta
has (we. Only female specimen available is headless.
Name derived from aglaes Cir, beautiful.
NEW SPECIIS OF STIGMODERA 143
Stigmaderg (Castiarina) atrs sp. nov,
FIGS IS 2K
Holotype, 4 kin W Pahima, Qld, 61.1986, A.
Sundholm, SAMA U 21192.
Allotype 9,4km W Paluma, Qld, 5.1986, & E
Adams, SAMA I 21193.
Paratypes: Od: 1 o, 1 9,4 km W Palumea, 41,1986,
A, Suyadhoim, ASSA: 1 o, same data as allotyne,
EAQA.
Colours Head black with blue reflections.
Antennas, segntents: 1, blue; 2-11, bronze-zreen.
Pronotun; and sculelluor black with bronze-preen
and blue reflections. Elvira yellow with following
black markings: basal margin; pre-medial fascia;
post-tnedial fasvia; apical mark, all confluent and
teaching margin enclosing eligi yellow spots, three
ln middle of each elytron and one on lateral margin
at cach humeral callus. Ventral surface bronze
green. Legs blue, Hairs silver.
Shape and sculpture: Head closely punctured,
median sulcus, medium length muzzle, Antennae,
segments: 1-3, obconic; 4-11, toothed, Pronorum
closely punctured, basal fovea extending to apical
margin as impressed line, basal notches represented
by glabrous area on each side more lateral than
medial; apical margin with slight medial projection,
basal margin bisinuate; laterally parallel-sided at
base, rounded to widest posteromedially, rounded
and narrowed to apex. Scutellum cordiform,
glabrous, cxcavate. Kagh elytron punctate-striate,
intervals convex, glabrous, 3rd, 5th and 7th faintly
raised at apex; laterally angled out from base,
founded at humeral callus, concave, rounded
posteriorly and narrowed to bispinose apex;
attenuated margittal spine, minute sutural spine,
miarein indented between, apices diverging, Ventral
surface with shallow punctures, edges of abdominal
segments glabrous, moderately hairy, hairs short.
Sy; truncate both sexes.
Size: Males, 9.4 ~ 3,2 mm (3). Females, LIL
3.8 mm {2}.
Male genitalia: (Fig. 1S), Sinuous in lateral profile,
apex curved upwards, Parameres parallel-sided
basally, rounded medially, straightened, widened,
Tounded apically, Median lobe pointed, sides angled
away. Apaphysis of basal piece moderately wide.
Remarks; Second member of 8 edligua species
group, In & obfigua Kerremans (Fig, IT), parameres
ate slightly widened basally, rounded at middle then
slightly narrowed, rounded to apex. Median lobe
blunt, sides angled away. Apophysis af basal piece
Sroad, The iwo species are easily distinguished as
S. oblique has green head, pronotum atid under-
surface and has yellow, black and red markings om
the elytra, S. efra could be confused with &
octosignata Carter which has similar markings,
That specics has: antennal segments 1-4, obconic;
bispinose elytral apices; marginal spine not atten-
uated to the same degree. Name refers to predomi-
nant colour and is derived from a@tra L., black.
Stigmodera (Castiarina) lauta sp. pov.
FIG, 25
Holotype 9, 4km W Paluma, Old, 4.(.1986, .4,
Sundholm, SAMA | 21194.
Paratype; 9, same data as holotype, ASSA-
Colour: Head black with blue muzzle, Antennac,
segments: 1, blue; 2-1], bronze-green. Pronorunt
and scutellum black. Elytra ycllow with confluent
black markings enclosing an elongate pre-medial
yellow mark on each elytron and yellow mark at
each humeral callus; red post-mnedial mark broader
Of margin, not reaching suture Veniral surface
bronze-green. Legs blue. Hairs silver,
Shape and sculpture: Head closely punctured,
plabrous, median suleus, medium length muzzie
Antennze, segments; 1-4, obconic; 5-11, toothed,
Pronotum shallowly punctured, glabrous, basal
fovea extending to near apical margin as glabrous
line; apical margin with slight medial projection,
basal margin bisinuate; laterally parallel-sided at
base, routed to widest part posteromedially,
rounded and narrowed to apex, Scutellum
cordiform, glabrous, excavate. Each elytron
punctate-striate, intervals Plat medially, convex at
base and apex, glabrous; laterally angled owt from
base, routided at humeral callus, concave, rounded
posteriorly, tapered to bispinose apex; marginal
spine larger than minute sutural spine, margin
rounded and indented between, apices diverging
slightly. Ventral surface with shallow punciures,
edges of abdominal segments glabrous, sparse short
hairs. Sy: leuncaté and indented medially in
females,
Size; Females, 10.3 » 3.8 mm (2),
Male genitalia; Unknown in this species. Male
genitalia of S gracilior Carter are illustrated
(Fig, IL) as only species {n group tor which males
are available, Parameres are curved upwards, spoon-
like apically, Parallcl-sided basally, widened,
rounded apically. Medial tobe pointed, sides angled
away, Apephysis of basal piece medium width.
Remarks: Fourth metnber of S. gracilier species
group on basis of external morphology. Easily
disunguished from other members of group by
elytral colour, S, yrereilier is red and black,
§. sulfured Deuduel is yellaw and black and &
ocfasignaia Carter is black with yellow spots.
Name is derived from fewzus L., splendid-
\44 S. BARKER
Stigmoder (Castiarina) gardnerae sp. nov.
FIGS IY, 2F
Holotype: of, between Black Hill and Swan Reach,
S. Aust., 6.x1L0984, S. Barker, H. barderwoude,
SAMA I 214195,
Allatype: 2, between Black Hill and Swan Reach,
S. Ause., 9x1. 1984, J & DB Gardner, SAMA | 21198,
Poratypes WAS 1 9, | 9, Afghan Rock,
Balladonia, |iii.1975, S Barker, SAMA: 2 ae,
Pindar, 21.1.1953, S. Barker, SAMA; 2 & ct, Borden,
27HiA9SH, .L AL. Watson, SAMA: | cr, Beverly,
Ho DuBoulay, SAMA; 2 oct, | 9, Bejugrding,
29.x11.1951, &. P McMillan, SAMA; | co,
Piawanning, 22.17.1950, R. RB McMillen, SAMA; 2
oo, Bucla, 64.1979, TL Hawkeswood, SAMA;
3 oo, Cranbrook, 4.1.1954, 4. ML Douglas,
WAMA; 1 cr, Yellowdine, 21.10.1962, L. McKenna,
A, M, Douglas, WAMA; 1 &, Lake Grace,
8/12:xi1. 1969, K. & E Carnaby, WAMA; 1 oo,
Fitzgerald Riv. N.P., 8/12.i1.1984, 23.11.1985, R. PR
McMillan, WAMA; 3 oc, Yellowdine, 14.1.1979,.
28.1.1979, T. M.S. Hanlon, WAMA;3 oo, 5 km
S Mt Holland, 27.11.1979, 7 Mf. 8. Henlon, WAMA;
Io, Dedari, 20,1,1982, & Jfantvch, 7. if Houston,
WAMA; | cr, Porresrfield, 10.11.1979, T Af, &.
Hanion, WAMA; | ot, Skim N Galena, 21.11.1979,
M. Powell, WAMA; 2 co, 1 9, Dedari, WAMA;
1 &, Cunderdin, Dec./Jan. 1919, WAMA; I cr,
Pixwanning, WAMA; Lo, Lake Grace, WAMA,
S. Aust bo, } OQ, Mundoora N.P, 31-xi1.1949, 8.
Barker, SAMA; 2. cot, McDonald-Ferries N.P.,
7.1.1970, 161.1980, S. Barker, SAMA; 3 ¢ or, same
dita as Holotype, IGAA; 1 9, between Black Hill
and Swan Reach, 2.xii. L984, J & D. Gardnes S,
Barker, IGAA; 1 @, Lyndoch, SAMA; 3 oe, 1
9, 5, Aust, SAMA; | or, no data, White coll.
SAMA, Vie. boo, Birchip, 10.1.1902, SAMA; 2
oor, Mallee clistricl, A. 7) Smith, NMVA; 1 &,
Merzinec, Jan. 1937, Fo, Wilson, NMVA; 1 co,
Gypsum, Nav. 1926, C. Qke, NMVA; 1 oO,
Inglewood, 14.1928, € Oke, NMVA; 2 cro,
Hattah, 25.x1.1950, J Plant, NMA; 1 cr, Sea Lake,
12.41.1916, Goudie, NMYA; | o, 1 @, Inglewood,
J Dixon, NMVA} 1 of, 1G, Mallee district, French
coll, NMVA; 2 oc, 2 9, Lille Desert, ¥,
Hateley, GBVA: G, Glemlee via Kiata, 8.ii1.1978,
G, G. Burns, GBVA; 1 @, 30,5 km NW Yanae,
15.41.1986, G. G. Burns, GBVA; | o&, Benetook,
25.41.1957, A. L. Brown, NMVA,
Colour: Head blue. Antennae, scgiments; 1, 2, blue;
3-11, bronze. Pronotum variably divided medial
blue mark, testaceous laterally, scutellum blue.
Elytra yellow with fullowing blue markings: narrow
{rasal marpin; clongate mark over humeral callus
mecting basal margin; small pre-rnedial spot on
eath side Closer to suture than margin; elongate
post-medial mark from margin 16 middle of each
elytron or reduced to a medial spot; diamond-
shaped pre-apical mark on Suture may or may not
reach apex, reduced to small spot in some
specimens; mark covering apex and spines, apical
margin red, Ventral surface testaceous. Legs: femora
and tibiae blue; tarsi bronze-gteen, Hairs. silver.
Shane and sculpture: Head closely punctured, broad
median sulcus, Antennac, seginents: 1-4, obconig;
5-1, toothed. Pronotum closely punctured, basal
fovea extending Lo centre a5 impressed line; apical
inargin straight, basal margin bisinwate; laterally
parallel-sided at base, angied outwards then
rounded at wides| part \% distance from base,
twpered) to apex, Scutellum scutiform, glabrous,
excavate, Each elytron punctate-striate, intervals
convex and punctured, the inner intervals shallowly
the outer intervals deeply; laterally angled out trom
base, rounded at humeral callus, concave, rounded
posteriorly, tapered to bispinose apex; very large
marginal spines, small, sharp sutural spines; margin
rounded and indented between, apices diverging,
margin subserrate from centre to apex, Ventral
surface with shallow punctures, edges of abdominal
segments glabrous, hairy, hairs short, 57; truncate
in both sexes, Legs, mule: tarsal pads absent on legs
2. and 3 on iarsal segments 1-3 replaced by median
spine.
Size: Males, 10.1 +010 » 3.740,05 mm (34).
Females, 10,7+0.21 » 4.04 0.09 mm (17),
Mate genitalia: (Tig. \V). Parameres gradually
widened, medially parallel-sided for short distance
then gradually rounded, abruptly rounded apically.
Median lobe pointed, sides angled away, Apophysis
of basal piece narrowed,
Remarks: Third member of $& mustelamajor
Thomson (Holotype male, Australia, MNHN)
specics group. 5, yrestetwmutyor occurs in castem
Australia li moderately high rainfall areas. Lateral
surfaces of pronotunk and ventral surface red,
elytral markings are similar lo S. gardnerae but
inuch heavier and it is much broader than &.
gordnene. S. erasma Carter (Holorype male,
Gippsland, NMVA) was synonymised with S&S
niustelamojor by Barker (1979). lt has heavy black
markings with elyiral villae and occurs in sub-alpine
eastern Australia. S. gerdnerae occurs in mallee
babitat and has lighter elytral markings than other
two. Genitalia of ull three are illustrated (Pigs. 1V,
TW, 1X), | consider all three valid species on hasis
of their male serlalia and dishunce distribution,
Named afier De J. Garciner, Adelaide.
NEW SPECIES OF STIGMODERA bs
Sligmodera (Castiarina) Jimturneri sp. nov,
FIGS ly, 2N
Holotype: &, Hill End, N.SW., 17.4,1984, FR.
Tiirner, SAMA f 21197.
Alletype: 9, Hill End, N.SW., 9.41,1983, ZR.
Turner, SAMA FT 2198,
Colour; Head, antennae, pronotum and scutellum
coppery with dark blue reflections. Elytra pale
yellow with following black markings with dark
blue and purple reflections: narrow basa! margin;
pre-medial fascia represented by elongate mark on
each side reaching margin but not suture and by
elongate mark slong siture, in allotype marks
confluent forming fascia angled anteriorly from
suture, reaching margin; broad post-medial fascia
reaching margin, projecting anteriorly and poster-
rorly along suture and anteriorly in centre on each
side from anterior margin; mark cowering apex;
marks may or may not be confluent along surure.
Ventral surface: sternum and legs coppery-red with
dark blue reflections; abdomen testaceous. Hairs
silver.
Shape and scalprure: Head closely punctured,
median sulcus, medium length muzzle, Antennae,
segments! 1-3, obconic; 4-11, toothed, Pronotum
closely pyinclured, basal fovea extending to centre
as glabrous line, basal notches on each side more
lateral than medial; apical margin projecting
medially, basal margin barely bisinuate; Iterally
angled outwards from base, rounded and slightly
bulbous posteromedially, tapered to apex. Scutelluin
sculiform, glabrous, flat. Bach elytron punctate-
striate, intervals convex, punctured, more so laterally
than medially; laterally angled out fron base,
rounded at humeral callus, concave, rounded
posteriorly and narrowed to bispinose apex; sharp
marginal sping, minute sutural spine, margin
rounded and indented berween, apices diverging,
apical margin subserrate. Ventral surface with
shallow punctures, edges of abdominal sezments
glabrous, moderately hairy, hairs moderately long.
Sy: truncate it male; rounded and shghtly pointed
in female.
Size: Male, 16.7 «x 6.6 mm [1). Female, 17.6 x
6.8 mim (1).
Male genitalia: (Fiz. 1¥). Parameres widened
basally, rounded and narrowed apically, Median
lobe with sharp point, sides rounded away until they
are horizontal, then dropping away vertically.
Apophysis of basal piece short, medium width.
Remarks; Member of S. fulviventris Macleay
species group. Aedeagus iol as heavily chilinised
as ceher members of §, fulviveniris Macleay species
group (Barker 1986, Figs 2), 2K, 2L). Closest to
&. arhertonensis, larger species. Distinguished from
other members by coppery-red colours of head,
thorax and legs, Named after Mr J, R, Tunes,
Hazelbrook.
Stigmodera (Castiarina) kanangara sp. nov.
FIGS 12, 2R
Holotype: Cc, Kanangara Walls, N.SW,, 20.71.1982,
JR. Turner, SAMA I 21199.
Paratypes: Z o ct, Hill End, NSW, 14/19,1,1984,
JR, Turner, TNA,
Colour: Head, antennae, pronotum and scutellum
green with gold reflections, Elytra orange with
following green or blue-green markings: narrow
basal margin; pre-medial fascia represented by
angled vitta over each humeral callus and elongate
mark along sulure; post-medial fascia reaching
margin, extending anteriorly and posteriorly along
suture or represented by medial mark on each side,
diamond-shaped mark on suture; pre-apical mark
on each side angled anteriorly and meeting at
suture; mark covering apex and spines or absent in
some specimens, murks may or may o01 be con-
Fluent along suture. Ventral surface: sternum and
all or part of 1st. abdominal segment green; at least
last four visible abdominal segments testaceous,
Leas green with gald reflections, Hairs silver,
Shape and sculpryre: Head closely punctured,
median sulcus, short muzzle. Antentiae, segments:
1-3, obgenic; 4-11, toothed, Pronotum closely
punctured, basal fovea extending tO centre as
glabrous line, basal notches on cach side more
lateral than medial; apical margin projecting
medially, basal margin bisinuale; laterally parallel-
sided at base, angled outwards, rounded to widest
posterobasally, rounded and narrowed to apex,
lateral margins compressed for more than half basal
length, Scutellum scutiform, elongate, glabrous,
flat. Each elytron punctale-striate, Intervals convex
and wrinkled; laterally angled out from base,
rounded at humeral callus, concave, rounded
posteriorly and narrowed ro bispinose apex; small
Marginal spinc shghily angled inwards, small! sutusal
spine, margin rounded and indented between, apices
diverging. Ventral surface with shallow punctures,
edges of abdominal segments glabrous, moderately
hairy, hairs medium length. S;: truncate in males,
Size: Males, 12.5 x 4.7 mm (3).
Male genitalia: (Fig, LZ), Parameres paralle-sided
basally, rounded outwards posteromedially,
rounded then parallel-sided, rounded off apically,
Median lobe pointed, sides obtusely angled away.
Apophwsis of basal piece medium width,
146 S. BARKER
Remarks: Member of S. straminea Macleay species
group (Barker 1986, Fig. 2N). Elongate body,
flattened pronotum and angle of marginal spine
distinguish it from other group members. Named
after type locality, an aboriginal place name.
Errata; Barker, S. (1986) Trans. R, Soc. 8, Aust, 110(1),
1-36, 30 May, 1986.
p, 2 insert [g] 8. cupricollis Saunders,
p. 9 paratypes of S. rufa sp. nov. 2 vo, 3 99,
Meekatharra, W.A., 5.vili.1972, not Mary Springs, W.A.,
8.ix.1970.
p. 20 paratypes of S. blackdownensis, 3 oo, 1 9,
Blackdown ‘Tableland, 23,ix,1973, E. E. Adams, ANIC,
AWHQ not EAQA, AWHQ.
p. 24 Allotype 9, WAMA not SAMA I 21172,
p, 33 insert pseuderythroplera Barker 1983, Trans, R. Soc.
S§. Aust. 107, 162 Q.
Acknowledgments
1 wish to thank the following for assistance: Dr
G. F, Gross and Dr E. G. Matthews, South
Australian Museum; Dr J. Lawrence and Mr T.
Weir, Division of Entomology, C.S.1.R.0.; Dr G. B.
Monteith, Queensland Museum; Dr T. F. Houston,
Western Australian Museum; Dr A. Neboiss and Mr
K. Walker, National Museum of Victoria; Miss
C. M. H. von Hayek, British Museum. (Natural
History), London; Dr P, Dessart and M., J. Cools,
Institut Royal des Sciences Naturelles de Belgique,
Brussels; M. A. Descarpentries, Muséum National
@Histoire Naturelle, Paris; Mr K. T. Richards,
Department of Agriculture, South Perth; Mr R. I.
Storey, Department of Primary Industry, Mareeba;
Miss M. Schneider, Department of Entomology,
University of Queensland; Dr D, Horning, Macleay
Museum, Sydney; Mr E, E, Adams, Edungalba; Mr
and Mrs G. G. Burns, Mornington; Mr & Mrs K.
Carnaby, Wilga; Mr H. Demarz, Quinn’s Rocks;
Mr T, M, S, Hanlon, Sydney; Mrs J, Harslett,
Amiens; Mr A. Hiller, Mt Glorious; Mr M, Powell,
Attadale; Mr. R. P. McMillan, Cottesloe; Mr J.
Sedlacek, Brisbane; Mr A. Sundholm, Sydney; Mr
J. R. Turner, Hazelbrook; Mr A. Walford-Huggins,
Molloy; Mr S. Watkins, Summer Hill; Mr G.
Williams, Lansdowne; Miss H. Vanderwoude, Dr
J. Gardner, Mr P. Kempster and Mr D. J. Williams,
Department of Zoology, University of Adelaide;
Australian Biological Resources Committee for
grants-in-aid of research.
References
BARKER, S. (1979) New specics and a catalogue of
Stigmodera (Castiarina) (Coleoptera: Buprestidae).
Trans. R. Soc. S. Aust. 103, 1-23.
(1980) New species and synonyms of Srigmodera
(Castiarina) (Coleoptera; Buprestidae). bid 104, I-7.
(1983) New synonyms and new species of Stig-
modera (Castiarina) (Coleoptera: Buprestidae). [bid
107, 139-169.
(1986) Stigmodera (Castiarinu) (Coleoptera:
Buprestidae); Taxonomy, new species and a checklist.
ibid 110, 1-36.
Wari, J. C. (1979) Abbreviations for entomological
collections. N.Z. Zool. 6, 519-520.
LACISTORHYNCHUS DOLLFUSI SP. NOV. (CESTODA:
TRYPANORHYNCHA) IN ELASMOBRANCH FISHES FROM
AUSTRALIAN AND NORTH AMERICAN COASTAL WATERS
BY IAN BEVERIDGE* & J, A. SAKANARIF
Summary
Lacistorhynchus dollfusi sp. Nov. is described from the spiral intestine of Mustelus antarcticus
Gunther, 1870 (type host), Pristiophorus cirratus (Latham, 1947), Galeorhinus australis
(Macleay, 1881) and Hypnos monopterygium (Shaw & Nodder, 1795) from Australian coastal
waters, and from Mustelus henlei (Gill, 1863) and Triakis semifasciata Girard, 1854 from
California. Metacestodes attributable to the species were identified from Synodus lucioceps
(Ayres, 1855) and Sebastes paucispinis Ayres, 1854, also from California. The new species is
distinguished by the presence of four characteristic bill-hooks on the base of the tentacle, and by the
distinctive shapes of hooks 1(11), 7(7') and 8(8') also at the base of the tentacle. The basal armature
of L. tenuis (van Beneden, 1858) is briefly described and illustrated and evidence provided that L.
tenuis may be a composite species. Cestodes from Californian elasmobranchs formerly assigned to
L. tenuis are re-identified as L. dollfusi, indicating a pan-Pacific distribution for the latter cestode.
KEY WORDS: Cestoda, Lacistorhynchidae, Lacistorhynchus, elasmobranchs.
LACISTORHYNCHUS DOLLFUSI SP. NOY. (CESTODA: TRYPANORHYNCHA) IN
i LASMOBRANCH FISHES FROM AUSTRALIAN AND NORTH AMERICAN COASTAL
WATERS
by IAN BEVERIDGE* & J, A. SAKANARIT
Summary
Beverinor, 1. & SAKANARI, JA. (1987) Lecistorhynchus dollfasi sp. nov, (Cestada: Trypanorliynchal in
elasmobranch fishes from Australian and North American coastal waters. 7rans. R. Soc. & Ans? 1163),
147-154, 31) November, (987.
Lacistorhynchus dollfusi-sp, nov. is described trom the spiral intestine of Mustelus antrcticus Gunther,
1870 (type best), Pristiozhorus cirretis (Latham. 1947), Galeorhinus australis (Macleay, 1881) and ff psros
nionopterygium (Shaw & Nodder, 1795) from Australian coastal waters, and from Mustelus henlet (Gill,
1863) and Triokis semifasciata Girard, 1854 from California. Metacestodes attributable ta the species were
iWentified from Syodus (vcioceps (Ayres, 1855) and Sebastes paucispinis Ayres, 1854, also trom Califarnia,
The new species is distinguished by the presence of four characteristic bill-hooks on the base of the lentacle,
und by the distinctive shapes of hooks 1(1'), 7(7'} and 8(8) also at the base of the tentacle, The basal armarnure
of . feruis (van Beneden, 1858) és briefly described and illustrated and evidence prowided that L. tennis
may be» vomposite species, Cestodes from Californian elasmobranchs formerly assigned to L. fenivis are:
reidentified as £. dollfest, indicaling a pan-Pucinec distribution for jhe latter cestode,
Key Worns: Cestoda, Lacistorhynchidae, Lueistorhyaciey, elasmobranchs.
Introduction
lacistarhynchus tenuis (van Beneden, 1858) is an
appdrently cosmopolitan cestode parasite of the
spiral intestine of sharks and rays. Dollfus (1942)
provided a detailed synunymy and deseription of
the parasire and summarized distribution records.
The type specimens were collected from
Galeorhinus gales (Linnacus, 1758) in the North
Sea, but the cestode is also known to occur in the
English Channel, olf the European and American
coasts of the North Atlantic, the African coasts af
the Atlantic and in the Mediterranean. Dollfus
(1969) subsequently provided additional morpho-
logical details based on specimens collected from
the Mediterranean. L, fenisis has been reported from
sharks and teleosts from the Pacific coasts of North
America (Young 1954a; Riser 1956; Yoge &
EFidmonds 1969; Pappas 1970; Mudry & Dailey 1971;
Buteau ef al, 1971; Heinz & Dailey 1974; Jensen ef
af, 1979, 1982) Moser ef af 1985) and South
America (Carvajal 1974; Duran & Oliva 1980;
Escalante & Carvajal 1984), Limited life cycle
Studies were carried out on material of Californian
origin by Young (1954b), Riser (1956) and Mudry
& Dailey (1971), while the life cycle was completed
experimentally by Sakanari & Moser (1985a) using
* Central Velerinary Laboratories, South Australian
Nepartment of Agriculture, c/o Institute of Medical and
Veterinary Science, Frome Road, Adelaide, 5S, Aust.
SUM). '
| Institute of Marine Sciences, University of California,
Santa Cruz, Catifarnia 95064, (15.4.
the copepod TJigrivopus califurnicus (Baker, 1912)
and the teleost Guerbusia affinis (Baird & Girard,
1853) as infermediute hosts, and Triawkis semifesciata
Girard, 1854 as the definitive host.
Studies on the effects of temperature and salinity
on life cycle stages (Sakanari & Moser 1985b) and
pathology induced by the plerocercoid in Merone
saxatilis (Walbaum, 1792) have also been carried out
(Moser ef af, 1984; Sakanari & Moser 1986),
The genus has nat heen reported from Australian
waters (Beumer ef al, 1982) but has been recorded
from Galeorkinus australis (Macleay, 1881) and
several species of teleosts From New Zealand
(Robinson 1959; Hewitt & Hine 1972), Recent col-
lections indicate that the genus ix not uncommon
in several shark species from South Australian
coastal walers including G. australis. In comparing
South Australian material with the description given
by Dollfus (1992), several discrepancies were noted
and tt became evident thar despite the detail of his
description, certain critical Features of the oncho-
taxy of £, fenuis have not been adequately
Jesenbed, Examination of specimens in the Dollfus
collection in Paris and material from Californian
hosts indicated that two distinet cestode species have
in the past been confused under the name L. feniis.
tn this paper, the Australian and Californian
material is described as a new species, and
additonal details of the morphology of L. ¢ensuts
are provided to enable differentiation of the rwo
species now considered to exist in the genus,
Fyidence is alsa provided indicating ihat L. ferwis,
as currently understood, may be a compuute of {wo
or more species
143 | BEVERIDGE & | A, SAKANARI
Materials and Methods
Cestodes collected from elasmobrauchs were
washed in sea water and fixed with bor 1%
buffered formalin ur bot 70% ethanol, In the
abserve of laboratory facilities (material collecied
on boats), spiral valves were Mooded wilh boiling
waler and a small yuantity of voncentrated formalin
was subsequently added. [In the labaratory, cestodes
were removed from the conrent, washed in water
aud stored in 70% ethiunol, Cestades were stained
With Celestine Blue, dehydrated, cleared im clove oil
aid mounted in balsam, Tentacles were removed
from some speclmens and were cleared tn glycerol,
then touted itv glycerine jelly,
Specimens of Lateistorkkpackus were borrowed
fron Me Srilish Museum (Natural History),
lendan (RMNH), the United States National
Museum Helminth Collection, Washington
(USNMHC}, the Dollfus collection from the
Muséum. national d'Histoive taturelle, Paris
(MNEIN), the Commonwealth Institute of
Parasilolagy, Ste Albans (CIP), and the Australian
Helminth Colleciion (AKC), South Australian
Museum, Adeluitle (SAM}, Unless athe'wise stated,
all Australian specimens have been deposited in
ARC, and American specimens in USNMHC.
Measurements are giver in che text in millimetres,
as the range of 10 individual measurements Followed
by the mean in parentheses, Terminology for che
morphology of trypanorhynch cestodes tallaws
Dollfus (1¥42) and Schmidl (1986), The hook uum-
bering system employed is that of Dullfus (1942),
Lacistorhvachos dollfisi sp. nov,
FIGS. t-t6
Tipes: Holorype, from spiral valve af Muste/tes
wntareticus Ginther, (870, Youne Rocks, south
cous of Kangaroo island, S. Aust,, 23.¥.7985, col-
lector B, G. Robertson, SAM V4UR5; 4 paraiypes,
same data, SAM V4084; 7 punilypes, same data,
AHC 32753; | paratype, sour coast of Kangaton
Island, S. Aust... 10.71.1985, collegjar B, G.
Robertson, USNMLIC 79544; 40 paratypes, same
data ALIC 42754; 3 paratypes, Goolwa, S% Aust,
21.1984, collector R. R. Martin, RMNIT 986.
9,29, 20-22; 19 paratypes, same data, AHC $2755.
Materia! excntinted: trom Mustelus antorericus Giiolher,
1370: types; fram Pristiephorus virratus (Latham, 1947):
Hi apcaimens, south voast Kangaroo Inand, S Ansa,
(AHL $2756); (rom Galewrhinus australis (Macteay, 1XX1):
3 specimens, Young Rocks, Kangaroo istand, 5. Aust.
(AIC $2757), S specimens, Pr Willunga, S. Aust.
(AH 312, $99); | specimen, worth-western Tasmania (CIP
D481), fram Notartivachyus cepedianis (Péeran, 1807);
2 specinens, Young Rocks, Kangaroo Island, S Aust,
(ANIC S2758), fram Hepes reomeplerveiuet (Shaw &
Nodder, 1795): | specimen, Holdtast Bay, & Aust, (AUC
52759), tron Mustelus testes (Gill, 1863;¢ 14 specimens,
Monterey Bay, California, U.S.A. (MNHN Bb28); |
specimen, Bodega Ray, Culitornia, USA. {MNHN Bb28);
trom THrdts sertifesciata Girard, 1854: 13 specimens,
Monterey Bay, California, U.S.A. (USNMHC 79591) from
Synodus ucinesps: 2 nelacestades, Matibu, California,
U.S.A, (USNMHC 74836); from Sebastes paucispinis
Ayres, 1854: | metacestode, Malibu, California, USA.
(USNMHC 76804),
Description (from types): Small cestodes, maximum
length 50, maximum width 1.20, maximum number
of proglottides 48, Scolex actaspedote, 1,52-2.40
(1-74) Jong, maximum width im pars bothridialis
0.26-0.41 (0.35) bothridia 2, patellitorm, with
slighlly indented posterior margin, shallow, without
Promineitt riet (Fig, 7}; diameter of bothridia 0.33,
0.38; pars bothridialis (.28-0.42 (0.36); pars vagin-
alis narrower than pars bothridialis, cxtremely
variable in length, 0.86-1.70 (1.12); pars bulbosa
slightly wider than purs vaginalis; tentacle sheaths
arranged in regular spirals; bulbs clorigate, 4-6 times
as long as wide, 0.33-0.46 (0.39) long by 0.06-O.1L
(0,09) wide; origin of retractor muscle im anterior
third of bulb, continued posteriorly by band of
nucleated, non-miuscular tissue (Fig. 13); scolex
markedly swollen immediately pusterior to bulbs,
swelling variable if length, stains intensely; bulbs
terminate either just anterior ro swelling or extend
info it; pars post-bulbosa, when present, 0.10-0.29
(0.16); scolex covered with prominent microtriches,
visible unuer light microscope,
Tentacles vecy lung and slender, up to 1.9 long,
0.20-0,40 (0.29) in diameter; slight swelling at base,
0.30-0.54 (0.41) in diameter: armature hetecomor.
phic, poeciloucanthous; armaiute at base of
proboscis distinctive: remainder relatively uniform:
hooks hallow. Base of probascis with bare area on
esiernal suface (Fig. 6); bothridial and anti-
bothridial surfaces al hase with 5 oblique rows of
filifonm huwks ranging in length from 0.006.0.014
(0.012) diminishing in size posteriorly (Fig, 4); ex-
ternal sui face of husal swelling with four pairs pf
large bill-hooks or four on antibothridial side and
three on bothridial side, cach with shurply recurved
blade, na guard and exiremely broad base; howks
af first pair (a) 0,022-0.026 (0.024) long, secund pair
(bh) 0,020-0.024 (0.021), third pair (c) O016-0.022
(0.019), Fourth pair (d) 0.08-0,014 (0.012) long; space
between pairs of bill-hooks on external surface af
proboscis occupied by six tilifurm hooks. Basal
hooks I{1')on internal surface (Fig. 4), represented
by three-four pairs of Jarge, slour. spinilorm hooks
followed anteriorly by four- five piiirs of overlappiie
hooks with small triangular blades, small guards
and narrow, clongate handles; anteriorly in meta-
basal region, hooks 11’) become lnrger, miere
Luencinatle O.012- 1,018 (1,014) long, with relatively
large blade and broad base: Basal honks 2(2'):
149
NEW CESTODE IN ELASMOBRANCH FISHES
2
SA ,
Pe oar BAA a
if” Que” : S : ae =<
A T NF [FF
j py Hel Lf fe
fi ELE )
s
£
a
LB
ra i
, ECS
Ye
ing
an
é eZ:
< os eer 7
ST an, Th La
ANsak SAMA AY
ae “SNS ere \
basal region,
; 6, basal Tegion,
+4,
gion, bothridial surface; 3, metabasal region, external surface
internal surface; bothridial surface to right; 5, basal region, bothridial surface; a-d, bill-hooks
external surface, bothridial surface to right; a-d, bill-hooks, Scale linc 0.01 mm. Hook numbering system follows
Dollfus (1942).
Figs 1-6, Lacistorhynchus dollfusi sp. nov. Tentacular armature. 1, metabasal region, internal surface 0.35 mm anterior
to base of tentacle; 2, metabasal re;
1) L BEVBRIDGE & 1. A. SARANARI
hooks of first four or five rows acicular, small:
anleriorly, nex five pairs adjacent to elongate
handle af hooks 1(1'), sinuous, apex usually directed
externally; in meta-basal region, hooks 2{2') become
large, Unelnate, 0.012-0,018 (0.015) long with
prominent hlade and extremely broad handle.
Hooks 3(3'), 44°), 3(5') similar in basal and meta-
basal regions, lung, slender, willy diminitive base,
handles absent, blade recurved at apex; lenpilis in
meta-hasal region; hooks 343’), O.018-0.022 (0.619)
long, hooks 4(4'), 0.018-0.020 (0,018) long; hooks
StS’), smaller, 0.4008-0.014 (0.011). Hooks 6{6') tiny,
acicular, slightly larger in basal region compared
with meta-basal revion, with thickened base, in-
seried in meti-basal region, slightly anferiar to level
of hooks 5(5') atid 747"), 6,003-0,006 (0.005) Tong
in Mmeta-lyasal region. Hooks 7(7) and 8(8')
commence anterior co Large bill-huoks; initially, 7
Jarger than 8, both with large, expanded bases and
acicular shafts: antetiorly, in meta-basal region, &
larger than 7, both uncinate with gently curving
blades, bases insignificant: hawks 7(7') 0.006-0.010
(0,008) tang; hooks 8(8') 0.010-0.018 (0.014) long,
Hooks 9(9), in form of double chainette, com-
menace anterior to level of bill-hooks as deltoid
plaques; hooks become smaller and acicular in
meta-basal region with slightly thickened bascs,
0.003 -0.0(K) (0,004) long.
Neck fegion extremely long and slender, 4.0-t4,0
(7.6), tegument with prominent folds, giving
appearance of segmentation; miture segnients
longer than wide, length 1.3-3.6 (2.3), width
O.O8-1.12 (1,97), acraspedote, Genital alnum in
posterior part of lateral margin, 0,9-2.2 (1.6) from
anterior end, surrounded by sphineter-like con-
densation of muscle fibres and parenchynia. Herma-
phroditic sac pyriform, |hin-walled, 0,32-0.40 (0,37)
by 0.63-0,19 (0,15); common genital duct of variahte
lenath, divides witht hermaphrodilic sac into
straight vagina and coiled cirrus, both surrounded
by glandular cells, Cirrus unarmed, gland cells react
arearest density in proximal region of cirries, prior
10 entry ito small, internal, crescentic semival
vesicle, External seminal vesicle absent; vas delerens
greatly coiled, passes slightly antenurly, then ails
posteriurly along mid-line to region of ovary; vasa
efferentia mot seen. Testes numerous, approximately
340 in number, occupying entire proglottis medulla.
Vawind pierces posterior wall of hermaphraditic sav,
passes towards ovary, Lerminaling in very slight
enkargement adjacen! to ovarian isthmus. Ovary bi-
lobed in dorsa-ventral view, gach lobe 0.20 0.40
(0.28) by 0.17-0.40 (0.24). Mehlis’ gland between
owartan Iohes, 0,14-0.21 (0.16) in diameter, Uterus
in midiine, extends almost to anterror extremity of
scament: yitellaria follicular, cireumecortical, follicles
0.016-G.032 (0,024) in diameter, L/rerine pore slit-
like, clearly visible in mature Segments, at level of
antetior end of uterus, Single gravid segment secnt
tree in gut lumen; 9.8 long by 2.2 wide; testes abserit,
ovarian lobes well developed, each lube 1.04 hy 0.48;
Mchiis’ gland 0.36 by 6.26; most of proglottis
occupied by uterus; uterus terminates below anterior
vextremily of segment; eggs ellipsoidal, opereulite,
0.046-0.050 (0.048) by 0.032-0,044 (0,033), aon-
embcyonated when laid,
Fiyorolegy The Tew species 1s named after R. Ph.
Dollfus in reeognition of his importance conrribu-
twins to Ihe systematics of the genus.
Lacistorhynckes jenyis (van Beneden, 1858)
FIGS 17-22
Material examined; rom Gatearhinus pateus (Linnaeus,
1758); 5 specimens, English Channel (BMNEH 1985.
17.17-20), | specimen, Scarhoroigh, England (BMNH
L979. 12.89.80); from Be/one befane (Lintaeus, 1758); 3
Metacestodes, S¢ie, France (MNHN Bb14; 55, 56, 58) 2
mctacestodes, Concarneau, France 4MNHN Bb); from
Mustelus canis (Mitchell, (815):9 specimens, Woods Hale,
USA, (determinal as Rayrchobuthruan bulbifer Dy
E, Linton) (USNMHC 4744, 4752, 7699); 7 specimens,
Woods Hole, U.S.A. (USNMHO 7950),
Description of tettacular armaiure: Meta-basal
armature similar to L, dolifust. Base of proboscis
with bare area on external surface; bothridial and
antibothridial surfaces with five oblique rows of fili-
form hooks, dimjpishing in size posteriorly; three
pairs of large bill-hoaks with sharply recurved
blades and extremely broad bases, Basal hooks 1(1")
On internal surface represented By three te four pairs
of stour spiniform hooks followed anteriorly by
lange Lriangular hooks with broadly curved blades
and diminitive handles, considerable discrepancy in
size between hooks of bulhridial and anti-bothridial
rows, Basal hooks 2(2’} clone, triangular to subu-
late, shehtly twisted. Hooks 3(3'), 4(4) and $(59
slender, with dimunitive bases, no handles, blade
recurved al tip, ciminish in size 3-5. Hooks 6(6')
tiny, acicular, Hooks 7(7') and &(8!) sub-triangutay,
large, With prominent, broad blade and expandedt
bases; base ul hook 8 overlain by blade af hook
7, Hooks 9(9') in form of double chainette, blades
tiny. bases swollen, Specimens from M, canix differ
ty shape oF hooks I(1') at base; greater disparity in
size between hooks of buthridial and ancibuthricial
rows than in European specimens; large hooks with
extremely clongate blades.
Discussion
The accouttt of the tisxonomie histary uf Lacy
forhvachus ienvis, compere list of synonyms, and
detailed account of its anatomy given by Dollfics
(1942), appareutly have been accepted by all subh-
Sequent authors, togerher with his lerntative con-
NEW CESTODE IN ELASMOBRANCH FISHES
/ h phe nigie + \ fe ¢
| f ee rs wig | D
[EGY HOOVES 6 | [pfs
{| eres eee on Efe iF 2
| bey QS OR ae
| | fl {> - 3 | te : Zc (on
\ ri oe 8 a ES:
| eagle We i
i lela® | i
ra)
et eRe i
ent,
te,
os
Figs 7-16. Lacistorkynchus dollfusi sp, nov. 7, scolex; 8, anterior region of strobila showing extremely long neck
region; 9, mature proglottis; 10, gravid proglottis; 11, neck region of cestode showing folding of surface tegument
giving impression of proglottisation; 12, tentacular bulb, showing insertion of retractor muscle; 13, detail of retractor
muscle insertion, showing continuation of retractor along internal wall of bulb by nucleated, non-muscular nysues:
14, female genital complex; 15, cirrus sac and genital atrium; 16, egg. Scale lines 0.1 mm.
1. BEVERIDGE & J, A. SAKANARI
i + 34
NEW CESTODE IN ELASNOERANCH FISHES \s3
clusion thal ihere Is a single, widely distributed
species in this gerius.
However, in exumining Australian specimens of
Lavistorkivnchas, certain discrepancies with Dollfus"
(1942) description were noted. Australian speciiviens
have four pairs of bill-hooks t= “serpe a bec" hooks
of Dollfus), or three pairs and a single extra book
on the external surface of the base of the tentacle,
yet Dollfus (£942) was not able to determine thy
exael nurnber of his speciinens. Furthermore, the
shape of hooks 11) on the base of the tentacle
clearly differed from those ilusurated by Dollfus
(1942, fig. 246). An examination of the specimens
upon which Dollfus had based his description and
his drawings (Pigs 244-254) showed that alrhough
the drawings were extremely accurate, the specimens
were very poor indeed, and che requisite morpho-
logical details were not available to him. However,
Mediterranean material in Dollfus’ collection 3g well
as British material from the type host Galeurhinus
galeus collected (rom (he North Sea indicated that
the Australian and Californian material described
above is different from £. fenwis. Because des-
criptions of the basal armature of L. tenuus are
incomplete and contain key distinguishing, features,
a setles of drawings is given (Figs 17-22), In f.
tenuis, there are three pairs of large bill-hooks on
the external base of the tentacle, compared with
four pairs or four and three hooks in Australian
specimens, Honks 1") of the basal region of L.
renuis are large and broadly curved with insignifi-
cant handles, as Showa by Dollfus (1942, fig. 244),
with a considerable disparity in size between hooks
in the bothridial and antibothridial rows, By con-
trast, tthe Australian specimens, hooks 1(1') have
a small blade and very long handles (Fig, 4), In
addition, hooks 7(7') and 8($’) of the basal region
are much more robust in L, tenuis than in Australian
specimens Because of these three major differences,
thy Australian specimens are considered Lo consti-
lute 4 species distinct From £. tenuis, Material
couispecitie with chat described above from Aus-
tralian hosts also occurs in elasmobranchs and
teleosts fram Californian coasts, Specimens of
Lacisterhynchus collected by N. W. Riser from
Mustelus henlei (syn. Rhinotriocts fentei) ia
Monterey Bay, California, and now im the Dollfus
collection (MNHN) as well asa further specimen
from the same host species fron: Bodega Bay,
California (MNHN 6128) and a series of specimens
from Triakis semifasciata from California are
identical with Australian specimens, as are metaces-
bigs 17-22, Lacistoriyachus tenuis (van Beneden, 1X58), Tentacular armvanie, specimens tron Afasre/us canis, Woods
todes collected (roi the leledsis Synwdus fucioceps
and Sebastes payeispinis both from Malibu,
Cahforma, listed in the material examined.
Bevause of the presence of the new species in
American waters, specimens of Rivrcoborthriuin
bulbifer Linton, 1889, identified by Linton, were
re-examined. This vestode species was described
from Mustelus canis from the north-eastern
Aulantic, and was placed by Dollfus (1942) as a
synonym of £ lertuis, Linton (1890) described a
further species, A, Aeterospine, which he timself
subsequently synonymised with AR. bulhtfer. No
material of R. Aeteraspine appears to be extane
(J, R. Lichtenfels. pers. comm,) and Linton’s own
synonymy of R. Hererospine with R. bulbifer has
therefore been accepted.
Matesial from the western Atlantic. from ibe
same lrost and localiry as Linton’s R. dulbifer,
differs From European material in having a greater
disparity in size between hooks | and VP at the base
of the tentacle, and a different hock shape, Owing
to the limited amount of material available [rom
European hosts, the extent of variation in oncho-
laxy is impossible to determine. The data available
at present sugvest that two species may be present,
and that &. Aubifer may represent an independent
species within Lacisforivachus. However, an
extensive collection of European specimens is re-
quired and the full extent of lntra-specific variation
dacumented hefore such decisions can he made,
Adequate collections do not exist {no European
museums to enable the problem to be pursued al
present,
Examination of material currently available sug-
gests that specimens of £ tenets from the
Mediterranean, the North Sea and the English
Channel are morphulogically similar, MacKenzie
(1985) has stupgested, an biological grounds, thal
more than ane species of Lacitarivrrias occurs
in fish in the Nonh Arlantic.
By contrast, the specimens of lacistariwnchus
from Australian and Californian elasmobranchs
clearly represent a species distinc! fron 2, fesitis
as presently defined and can be readily distin-
guished by features of the onchotaxy,
“L. tenuis” as repurted by Riser (1956) and Jensen
et at, (1979) Tram California is new attributed to
L. dollfusi, Likewise, the life-evele studies of
Sakahari & Moser (1985 a, 6), Moser ef af, (1984)
and Sakanati & Moser (1984) are alsn atrribintedd lo
Z. dotlfun vather than 2. remus. to view of the
findings reported here, all other western American
Hole, U.S.A, 17, basal region, internal surface: 18, basal region, external surface; 19, basal region, borhridial surface.
Tentacular armature, specimens from Belone helane, Séte, France; 20, basal pevion, internal surface; 21, basal region,
external slirface) 22, basal region, borhrtdial surtacc. Seale tines 0.0) roam, Hook qumbering vystem follows Dollfus
(1942); a-c, bill-hooks
154 | BEVERIDGE & J. A. SAKANARI
records should be verified betore the occurrence of
7, fennis in the Pavifie region is accepted.
Acknuwledgments
Thanks are duc to BG. Robertson, M. G.
O'Callaghan and R. R. Martin for collecting clas
mobranchs and/or cestndes; to Drs R. Bray, I R,
Lichtenfels, A, Petter and A. Jones for the loan of
specimens; and to the Australian Biological
Resources Survey for financial support.
References
Heimer, J. P., ASHBURNER, L. D. Burpury, M. E.,
Jette, R. & IaATHAM, D. J. (1982)-A checklist of the
parasites of fishes {rom Australia and its adjacent
Antarctic territories. Technical Cotamunication No. 48
ar the Commonwealth Institute of Parasitology.
Commonwealth Agriculiural Bureaux, Varnham Royal,
England,
Bureau, G. H., Simmons, J. &, Bracu, D. EL, Hotz,
GG. & StkeMan VOW 1071) The linids-of eesrodes
from Pacific and Atlantic coust Triakid sharks. 2
Parasitol, 57, 1272-127K,
CARVATAL, J. (1974) Records of cesiodes from Chilean
sharks. Ibid. 60, 29-34.
Doivr.ts, R. P. (1942) Etudes critiques sur les
Tétrarhynques du Muséum de Paris. 47ch. Mfus. nett.
Hist. nat, Paris, 6 ene ser WW, 1-466,
——{1969) De quelques cestodes téirarhynques
(hétéracanthes er pécilacanthes) recoltés chez des
poissons de la Méditerranée, Viv et Milieu 20, 491-542,
Duran, L. B. & Oxiva, M. (1980) Estudio parusitologica
en Merluccius gayi peruanus Gingsburg, 1954. Bol.
Chile. Parasit, 35, 18-21,
ESCALENTE, FL. ak CARVAJAL, J. (1984) Larval
trypunorhynch vestodes from Peruvian teleost fishes,
with descriptions af two new species, Siud. Nearrop,
Fauna Erty. 19, 185-194,
Herz, M. L. & Daitey, M. D. (1974) The
Trypanorhyncha (Cestoda) of elasmobranch fishes from
southerft California and northern Mexivo, Prod, Hel,
Soc. Wash. 41, 161-169.
Hewitt, G. C. & Hise, P.M. (1972) Checklist of
Parasites of New Zealand fish and-af their hosts, MZif
Mar, Freshw. Sci. 6, 69-114.
Junsen, L. A., Moser, M. & Heckmann, R, A. (1979)
The parasites of the California lizardlish, Synodus
luctoceps, Prac. Helm. Sog. Wash, 46, 281-284.
. & Dairy, M. 0 (1982) Parasites of
Rocaccio, Sehastes paucispinis, trom southern and
ventral California, /bie/. 49, 314-317,
Linton, E. (1890) Notes on eritozou of murine fishes of
New England. Part I. Annual Report of the
Commissioner af ish and Pkishertes’ for 1887,
Mashinglen, 719-900,
Machemzin, K. (1985) The use of parasiies as biological
tags in population studies of herring (Clipe
hureneus dyin the Nerth Sea and to the nowh ane west
of Scolanad. J cons. Jat. expl mer. 42, 33-64.
Mose, M., SAKANARL, LL, WELLINGS, S. & LINDSTROM,
K. (1984) tncornpatibiliry benween San Francisco striped
bass, Morone sexatilis (Walbaum) and the metacestode,
Lacistorhynchus tenuis (Beneden, 1858). J. Fish Diy.
7, 397-400,
—----, , REitty, C. A. & WHIPPLE, J. (1985)
Prevalence, intensity, longevity and persistence of
Anisakis. sp, Sarvae and Lacissorhynehus tenisis
inelavestodes in San Francisco striped bass, National
Oceanic and Atmospheric Administration, National
Marine isheries Service, Technical Report 29, 1-4,
Munry, 2 R, & Dattry, MLD. (1971) Postenbryonic
development of certain tetraphyllidean and
trypanorhynchan cestodes. with a possible alternative
lite cycle for rhe order Trypatiorhyneha, Can. £ Zoul.
49, 1249-1253,
Paveas, P. W, (1970) The trypanorhynchid cestodes from
Humboldt Bay and Pacific Ocean sharks. J. Parasitol,
56, 1034,
Riser, N. W. (1956) Early larval stages of two. céstodes
from clasmobranch fishes. Proc. Helm. Sac. Wask, 23,
120-124,
Romnson,, E.'S. (1959) Records of cestodes from marine
shes of New Zealand, Trans, R. Soc, N.Z, 86, 143-153.
SAKANARI, J. & Moser, M. (1985a) Iniectivily of, and
laboratory infection with, an elasmobranch cestode,
Lacistorhynchus tenuis {Van Benederi, 1858),
Parasitol, 71, 788-791,
——;; (1985b) Salinity and temperature effects on
the eges, coracidia, and procercoids of Lacistorhynchs
tenuis (Cestoda: Trypanorhyncha) and induecd
mortality in a first intermediate host. [bid. 71, 583-587-
——&{———+(1986) Lesion induction by the
plerocercoid Lacistorhyachus tenuis (Cesioda) and
wound healing in the striped bass, Morone saxarilic
(Walbaum). J. Fish Biol. 28, 289-296,
ScHMIDT, G. D. (1986) Handbuok of Tapeworm
Identification, (CRC Press {nc., Boca Raton, Ploriday
Younc, R. T. (1954a) Cestodes of sharks and rays yn
southern California. Proc, Helen Soc. Wash. 21,
106-112,
(1954b) A nate on the life eyele of Lacistorkynoties
tenuis (van Beneden, 1858), a cestade of the leaparel
shark, /bid. 21, 2.
Voor, M. & Epmenos, [L. (1969) Matching in vitro of
corucidia from oncospheres of Lacutorhpachys tenuis
(Cestoda: Tetrarhynchidea). J. Parasitol, 55, 571-573,
A MEGAFOSSIL FLORA FROM THE EOCENE OF GOLDEN GROVE,
SOUTH AUSTRALIA
BY D. C. CHRISTOPHEL & D. R. GREENWOOD*
Summary
A new Eocene plant megafossil locality is reported from near Golden Grove, South Australia. Well
preserved mummified leaves, fruits and flowers and impressions are abundant in a lens structure
within the East Yatala Sand Pit operated by Monier. Preliminary analysis has identified a rainforest
flora containing Elaeocarpaceae (Sloanea/Elaeocarpus), Lauraceae, Myrtaceae (Myrtaciphyllum),
Proteaceae (Banksieaephyllum, aff. Neorites), Sterculiaceae (Brachychiton), Podocarpaceae
(Decussocarpus, Podocarpus), and the fern Lygodium. Physiognomic analysis of the leaves
suggests that the closest analogue is with Complex Notophyll Vine Forest. The Monier Golden
Grove Eocene flora has some taxonomic similarity in common with other southern Australian
Eocene floras, but has a distinctive character which supports the idea of a regionally diversified
rainforest vegetation in southern Australia in the Eocene.
KEY WORDS: Fossil, Eocene, Golden Grove, South Australia.
A MEGAFOSSIL FLORA FROM THE EOCENE OF GOLDEN GROVE,
SOUTH AUSTRALIA
by D. C. CHRISTOPHEL & D. R. GREENWooD*
Summary
CuHerisToPHEL. D.C. & Greenwoop, D. R.. (1987) A megafossil flora from the Eocene of Golden Grove,
South Australia, Trans, R. Soc. S, Aust. 11403), 155-162, 30 November, 1987.
A new Eocene plant megafossil locality is reported from near Golden Grove, South Australia. Well
preserved mummified leaves, fruits and flowers and impressions are abundant in @ lens structure within
the East Yatala Sand Pit operated by Monier. Preliminary analysis has identified a rainforest flora containing
Flaeocarpaceae (Sloanea/Elaeocarpus), Lauraceae, Myrtaceae (Myrtaciphyllum), Proteaceae
(Banksieaephyllum, aff, Neorites), Sterculiaceae (Brachychifon), Podlocarpaceae (Decussocarpus,
Podocarpus), and the fern Lygodium. Physiognomic analysis of the leaves suggests that the closest analogue
is with Complex Notaphyll Vine Forest. The Monier Golden Grove Eocene flora has some taxonomic
similarity in common with other southern Australian Eocene floras, but has a distinctive character which
supports the idea of a regionally diversified rainforest vegetation in southern Australia in the Eocene,
Key Worps: Fossil, Eocene, Golden Grove, South Australia,
Introduction
In. January 1986, a fossiliferous clay lens was
uncovered during sand quarrying operations at the
Monier East Yatala Sand Pit in Golden Grove,
South Australia (138°43'30'., 34°47'10'') located
approximately 25 km north-east of Adelaide city
centre (Fig, 1), The lens was found amidst cross
bedded, fresh water sands and the entire
depositional sequence suggests a meandering stream
system with possible lacustrine elements.
While this is the first megafossil flora described
from the Golden Grove area, other plant fossils have
been reported from the region,
Eocene clays containing dispersed plant fossils
have been described in several papers (e.g. Lange
1978a, 1978b, 1980), but a precise description of the
location of the deposit is not given in any of them,
As a recent drilling program by the S, Aust, Mines
and Energy Department has indicated that the
lateral extent of the new lens is extensive (Neville
Alley pers comm.) it is possible that these earlier
reports Were based on material from the same
system, However, it is known that the earlier
material did not come from the Monier quarry, and
hence precise geological comparison between the
earlier reported material and the Monier Golden
Grove Locality may never be possible, They may be
palynologically correlated. however.
A preliminary palynological analysis (W. K.
Harris, Western Mining Co, pers, comm.) placed
the lens at the base of the Proteacidites pachypolis
Zone of the Eocene. A second palynological
analysis (Neville Alley pers, comm.) supports
placement in this zone. The deposit is therefore
“ Department of Botany, University of Adelaide, G.P.O.
Box 498, Adelaide, S. Aust. SOOL.
either just younger than the Maslin Bay deposit
located 30 km south of Adelaide or equivalent to
it, The Monier Golden Grove deposit is slightly
older than the Anglesea deposit in Victoria
(Christophel, Harris & Syber 1987) (Fig. 1).
f
, “
\h GOLDEN GMOVE PA
MADLIN BAY @ ,NEMLLGA
Pu,
wuoLRSEe Sy
Fig. | Map of eastern Australia giving the location of the
Golden Grove locality relative to the Eocene localities
of Maslin Bay, Anglesea and Nerriga, and lwo of the
sites trom. which litter was collected for the
physiognomic comparison,
156
Materials and Methods
Portions of the lens are highly oxidised and plant
remains are preserved as brown impressions in beige
matrix with no organic remains present, The
majority of the lens contains carbonized remains
within a grey matrix, and mummitied leaves are
floated from this material by placing blocks of the
clay in a 2-3% hot aqueous solution of hydrogen
peroxide to disaggregate the matrix, Many of these
leaves are naturally translucent and can be photo-
graphed directly with transmitted light (Figs 2-6).
The rest are black, opaque, and were observed and
photographed with UV light to study the venation
detail using the technique of Christophel ef al.
(1987). This type of preservation is shown in
Figs 7-9,
A sample cuticle is then prepared using the
method discussed in Christophel ef a/, (1987) and
the leaf is mounted between glass slides in Phenol
Olycerin. Jelly.
All leaves illustrated in this paper are housed in
the Adelaide University Palaeobotany collection.
Fruits and flowers obtained from the macerate are
stored in 1% phenol in 50% ethanol in sealed vials,
in the same collection.
¥ 4
an
if
~~
a
Figs 2-4. Transparent mummified leaves from Golden Grave. Scale bars =
Sloanea/Elagocarpus; Figs 3., 4, Lauraceae,
B.C. CHRISTOPHEL & D. R. GREENWOOD
Components of the Megafossil Flora
The commonest leaf taxon from the munsmilied
horizons at Goldeti Grove is a serrate-margined
notophyll (Fig. 2), This taxon has superficial
similarities to Sloanea/Elaeecarpus (Elaeocar-
paceae). A recent study of extant species of this
family found that leaves of Elaeocarpus L. and
Sloanea L. could not be separated, but that they
consistently clustered distinctly from other genera
in the family and from other architecturally similar
families (Moira Turnbull Adelaide University pers,
comm.). The Golden Grove fossils similar to Fig. 2
consistently clustered with the extant Sloarnea/
Elaeocarpus complex.
Aiother common taxon in the Monier Golden
Grove flora is Banksieagephyllum Cookson &
Duigan (Fig. 9), As the name suggests, the leaf is
very similar to those found in the tribe Banksieac
(Proteaceae), but as Christophel & Greenwood
(1987) suggested, this could also represent the foliar
organ of Musgraveinanthus Christophel (1984).
This is supported by the occurrence of
Musgraveinanthus in the Monier Golden Grove
deposit, which is the only locality other than the
type locality (Anglesea) from which it has been
\ cm. Fig. 2, Blaeocarpaceae aff,
GOLDEN GROVE FOSSIL FLORA is?
reported. Banksieueidites arcuatus Stover, the pallen
iype found in Musgraveinanthus inflorescences, is
also found as a dispersed grain at Galden Grave.
The Lauraceae has been suggested as an
important family in Australian Barly Tertiary floras,
Based on the characters discussed by Hill (1986) as
diagnostic of the family, we haye confirmed at least
two comman leaf taxa at Golden Grove (Figs 3, 4)
which may be assigned to that family. The
importance of the family in the Monier Golden
Grove flora is therefore confirmed.
Leaves of Myrtaciphyl/um Christophel & Lys are
also frequently encountered in the Golden Grove
flora (Fig. 7). However, a preliminary investigation
indicates that the Golden Grove Myrtaceae leaves
belong to different species than the two described
by Christophel & Lys (1986),
‘Two leaf types collected at the Momier Golden
Grove locality are known only rarely from other
Early Tertiary deposits, Table t. These are leaves of
Brachychiton (Sterculiaceae) and the fern Lygodium
(Schizeaeaceae). Brachychiton Scott & Endl. is very
common as impressions in the leached portion of
the Jens, where it is recovered in both a three and
five lobed form. Less common in the mummitied
portions of the lens, it has thus far only been
collected in a three-lobed form from that material
(Fig. 8). Lygodinum is not only recovered as sterile
pinnules (Fig, 6) but has alsa been collected as
numerous fertile fronds, These can be seen to con-
tain spores, which have yet to be studied and com-
pared to their extant counterparts.
The gymnosperms are represented at Golden
Grove by two genera of the Podocarpaceae,
Decussocarpys Laubenf. and Podocarpus L’Herit.
ex Pers. sens. strict, Decussocarpus twigs are
common, and appear morphologically intermediate
helween D. maeslinensis Blackburn, described from
Maslin Bay, S. Aust., and D. brownes (Selling)
Greetiwoad, from Anglesea (Greenwood 1987), It
is. quite likely that the three therefore represent a
range of forms in one highly variable taxon, Similar
variation can be seen in modern examples of the
genus [eg. D. vitiensis (Seeman) Laubent.,
Greenwood 1987], Golden Grove material of this
species has been illustrated earlier by Christophe!
& Greenwood (1988),
Podocarpus leaves are less common, and while
much longer, have epidermal features: identical to
Podocarpus platyphyllum Greenwood described
from Anglesea (Greetiwood 1987),
One further leaf type is reasonably common and
has not been reported previously. These leaves are
pinnately compound (Fig. 5} and on the basis of
their stomatal arrangement and epidermal hair
bases can be ascribed to the Proteaceae. They bear
a superficial similarity to immature leaves of
Neorites L.S,Sm,
While the identity of no other leaf taxa have been
confirmed, a preliminary estimate of approximately
30-35 leaf types for the deposit is made, However,
the present collection contains only approximately
400 leaves, and so the preliminary nature of the
estimate must be emphasized, Four fruit/flower
types have been thus far recovered, but only
specimens of Musgraveinanthus have been
identified.
Physiognomic Analysis
The general elliptic shape, broad lamina, and the
presence of well developed ‘drip tips’ on many leaves
(ee. Fig. 4), strongly suggest that the Golden Grove
flora was derived predominantly from rainforest
vegetation. These features are often cited as being
characteristic of rainforest leaves, particularly
lowland tropical rainforest (Richards 1957). The
presence of typical rainforest taxa eg. Elacocar-
paceae (S/oanea/Elaeocarpus), Proteaceae
(Musgraveinae, Neorites), Myrtaceae (Syzyedum
Tae 1, Megafossil taxa at Australian Localities,
Loéalily Galden Grove
Taxa
Maslin Bay Anglesea Nerriga
_— FT SSS
Austrodiospyros
Museroveinanthus
Elaeocarpacese
Myrteviphyllum
Brachychiton
Devuxsocarpus
Podocarpus
Gvrinosianiu
Zamiaceac
Lauraceae
Lygodiurnt
Banksleaephyllum
Diversity
xONNOAY
AAD
30-35
R
?
be
FAGAN ANBKORT
' 10K) +
-ee SE
C — common (> 10%); X = present (1-10% — possibly more common, but nat assessed); R = rare (< 1% of
flaray; ? = similar bul unconfirnied.
158 D. C. CHRISTOPHEL & D, R. GREENWOOD
Figs 5, 6. Transparent mummified leaves from Golden Grove. Scale bars = | cm. Fig. 5. Proteaceae aff. Neorites;
Fig. 6. Fern pinnule, Lygodium.
Figs 7-9. Opaque mummified leaves photographed using
UV light source. Scale bars = 1 cm. Fig. 7.
Myrtaciphyllum; Fig. 8. Brachychiton; Fig. 9.
Banksieaephyllum.
type), Lauraceae, Podocarpaceae (Decussocarpus,
Podocarpus) strengthens this impression, although
representatives of some of these families are also
found in more open forest types. It is appropriate
therefore to compare the foliar physiognomy of the
Golden Grove flora to that of modern rainforest.
The use of foliar physiognomic analysis for the
interpretation of fossil leaf-beds based on forest
canopy data has been criticized for ignoring the
effect of transportation and the differential input
and preservation of leaves prior to sedimentation
(Martin 1986; Christophel & Greenwood 1988).
Recent research, however, suggests strongly that
leaf-litter from each of the main rainforest types
recognized by Webb (1959, 1968) and Tracey (1982)
has a unique physiognomic “signature” and that this
signature is often retained in stream-deposited leaf-
litter (Christophel & Greenwood 1988). Work in
progress by the second author supports this. The
physiognomic features of primary importance are
length, width, shape and margin type. Leaf length
only will be considered in this paper, as it is
sufficient to illustrate the physiognomic signature.
Fig. 10 shows the frequency distribution of leaf-
length for the four main rainforest types recognised
by Webb (1959, 1968) and the frequency distribution
GOLDEN GROVE FOSSIL FLORA 159
for the Golden Grove flora using mummified leaves.
As can be seen from the figure, the extant rainforest
types can be clearly separated on leaf-length alone,
Fig. 11 shows the same data plotted as a cumulative
percentage for each length class. The CNVF
(Complex Notophyll Vine Forest) forest type has
been presented as two separate sites to better
illustrate the intrinsic physiognomic variability of
litter from NE Queensland (The Crater) and N.SW.
255 + +
NM CMVF CNVF
* nai2 n=mi2
+. +
205 « +
~~ ~
~~ +
te +o
te >
155+ « +.
> al > st
+ we ~ *
= eH - .
aa Lal - ~
105 ~ 7” - aa
Lal -— ee
nH - Hea
He <= 4
. oom) te
55 s + ae
s >
~~
7+ + + + =a tHe PS
5 10 15 20 25 5 10 15 20
255 +
tT GOLDEN GROVE
Leaf Length 155
( 10 mm classes )
'
5 10 15 20 2
(Dorrigo) from this type. Standard errors for the
data from these sites are presented in Table 2. These
results indicate that the Golden Grove Eocene
vegetation was physiognomically closest to CNVEF
from The Crater N.P. and less so to CNVF from
Dorrigo in northern N.SW. Dorrigo and The Crater
fall within the Mesotherm seasonal Torresian zone
of Nix’s bioclimatic classification of Australia (Nix
1982).
I
f snve t MMF
i n=12 | ned
+
: T
= +
D di
; 4
e +
Fi 4
. =
-_ oo
Land i
ee wa
HH Lal
eo _
Ld ee
Lal
= + + + +—
5 10 15 20 25 5 10 15 20 2
ne 5
|
|
ore
a a a
5 10 15 20 25
Frequency %
Fig. JO. Frequency distributions of leaf length (10 mm classes) for Microphyll Mossy Forest (MMF), Simple Notophyll
Vine Forest (SNVF), Complex Notophyll Vine Forest (CNVF), Complex Mesophyll Vine Forest (CMVF), Golden
Grove, and Anglesea. Forest nomenclature follows Webb (1959, 1968) and Tracey (1982). Each of the data sets
for the modern forests is based on four samples of 200 leaves collected from litter (Christophe! & Greenwood
1988). Golden Grove data are based on 156 complete leaves from the initial 1986 collections, Standard Errors
are plotted.
16D 19, ©. CHRISTOPHEL & D. R. GREENWOOD
Comparison with other Eocene Floras
As mentioned above, the Monier Golden Grove
flora is similar in age to the Maslin Bay flora and
is somicwhat older than the Eocene Anglesea flora.
It is younger than the Nerriga flora (Hill 1982) but
can still be usefully compared with it, as all are
Eocene, Table | Compares the major (as well as. some
of the unique minor) taxa from the four deposits,
Perhaps the strongest similarities ta the Golden
Grove megafossil flora may be seen in the Anglesea
locality. While the diversity of the Anglesea Mora
is shown as much higher (100+), this represents a
composite of six separate fossiliferous lenses. Taken
separately, the lenses at Anglesea have a diversity
much more similar to Golden Grove (Christophel
et al. 1987). The strongest links between the two
are the very similar Elaeocarpaceae leaves,
Musgravelnanthus, Myrtaciphyllum and
Brachychiton, They also have in common
Decussecarpus, Podocarpus, Lauraceae,
Banksiewephyllum and Lygodium, but these are not
restricted to thase localities. Physiognomically,
however, Golden Grove is interpreted as CNVF,
whereas the New Site lenses at Anglesea were
interpreted by Christophel (1981) and Christophel
& Greenwood (1988) as being closest to SNVF
{Simple Notophyll Vine Forest) from N.E.
Queensland (Fig. 11).
The diversity of Golden Grave is similar to
Netriga, but there are very few shared taxa, and the
only feature in common is really the shared
importance of the Lauraceae, Maslin Bay has a few
taxa in common with Golden Grove, but in general
Maslin Bay has a high diversity with no truly
common taxa, while Golden Grove has fewer, well
represented taxa and hence suggests a quite different.
floristic association.. Preliminary physiognomic
analysis. of the Maslin Bay flora by the second
author has detected minor differences which suggest
that the Maslin Bay flora was sourced from a
warmer vegetation type (Greenwood unpubl.).
The absence of both the Zamiaceae und
Gymnostoma Johnson. at the Golden Grove locality
is surprising considering the commonness of both
at Anglesea and their presence in the other deposits.
While further collecting may reveal either or both
of these taxa, their absence to date may be taken
as evidence that they are not major components of
the Monier Golden Grove flora.
The preliminary investigation of the Golden
Grove Nora has revealed a vegetation typical of the
southern Australian Eocene floras studied to date,
in that it appears to have been sourced from a
rainforest vegetation (Complex Notophyll Vine
Forest sensu’ Webb, 1959, 1968) of reasonable
diversity in a mesotherm environment (sensu Nix
1982). There are differences between it and the other
documented floras of similar age, as is expected
based on the temporal and spatial differences in the
localities. While numerous similarities in taxa
present have been highlighted in Table !, it must
be emphasized that these are similarities in natural
and form genera, and there is no evidence yet
(possible excluding Podocarpaceae) to suggest that
Tapbe 2, Standard Error Values for Curiulative
Frequency of Length for Fig. 11.
length class (mm) MMFTt SNVFY
e's
HER ARCOLA ROmW Eins
an
wm
PSH rH NNYMSS
Mpa kowe ah
SSSSCOSH SK Kew aDwMS
195
205
215
225
4
>
Pll} ide bi tb
Ss
Dorriga* ‘The Cratec* CMVF*
_ — 0.1
0.2 1.3 24
a9 3.7 3.4
0.7 3.1 3.2
0.8 1.4 24
1.5 1.8 2.3
4Q 1:2 3.1
3,9 1.1 at
23 00 4,2
1.7 0.9 3.7
(.3 1.1 4.2
0.9 5 29
0.6 1.0 1.8
0.4 0.8 Lt
02 0.6 0.7
0,2 0.5 0.5
0.2 0.2 0.4
0,3 — 03
0,1 — o4
0.1 _— 0.2
- — 0.)
*a - 4samples of 224 leaves,
fn — 4 samples.of 224 leaves lor 3 wiles.
GOLDEN GROVE FOSSIL FLORA
100 rt s
a
.
90 bd
s e
BO ° 3
7
# 70 . °
a 7 ¥
a 60
z .
a +
cx 50 ‘
o s
>
we 40 q *
S
E °
6 30 ,
e
20 . 7 .
j
{ *
io! é }
: 5g AY
25 35 45 55 65 75 BS 95
105 116 125 135 145 155 165 175 185
14)
MMF <NSW)
SNVF (NEQ)
CNVF (Derriga)
CMVE (NEG)
CNVF (The Crater)
Golden Grave
Js 6¢ < @ @
9
=a ——<$_——___~
195 205 215
Leaf Length (10 mm classes)
Fig. 11, Cumulative trequency histograms for the data presented in Fig. 10. Two separate localities for CNVF are
given: the Crater National Park (open triangles) and Dorrigo (solid triangles). These two localities reflect the extremes
of the range seen in litter samples from N.E. Queensland and N.S.W.
the same species are occurring in the different
localities,
The Monier Golden Grove flora represents South
Australia’s only known mummified leaf flora still
available for collection and study (a collection of
several thousand Maslins Bay specimens exists in
the Botany Department af Adelaide University, but
the locality is no Jonger collectable), and it is hoped
that further work will provide us with deeper
insights into the evolution of South Australia’s flora,
Much of South Australia’s coal reserye which. is
currently being considered for development is
Eovene in age, and hence Golden Grove will
hopefully provide a benchmark for studies of these
floras when they are eventually developed.
Acknowledgments
The authors wish to acknowledge the help and
cooperation of David Keane, Rex Rowley and
Maurice Burton of Monier Ltd. We are also grateful
to Neville Alley (S.A. Mines and Bnergy
Department) and Neville Pledge (S.A, Museum) tor
bringing the deposit to our attention, Research on
this locality is supported by a joint grant from the
S.A. Mines & Energy Department, S.A, Department
of the Environment and Planning, and Monier Ltd.
The plates were prepared by Susan M,. Shaw,
References
CueisToPpHEL, D. C. (1981) Tertiary megafossil floras of
Australia .as indicators of floristic associations and
palaeoclimate. Ch. 12, Jn Keast, A. (Kd.) “Ecological
Biogeography of Australia”. pp. 379-390, (Junk, The
Hague.)
(1984) Early Tertiary Proteaceae: The First Floral
Evidence for the Musgravetnac. Aust J Bol. 32,
177-186.
, GREENWOOR, D, R. (1988) A comparison of
Australian Tropical Rainforest and Tertiary Fossil Leaf
Beds. Proc. Ecol, Soe. Aust. 15.
: Harris, W. K., & Syecr, A. K. (1987) The
Eocene flora of the Anglesea Locality, Victoria.
Alcheringa 11, 403-324,
, & Lys, 5. (1986) Mummified Leaves of Two New
Species of Myrtaceae from the Kocene of Victoria,
Australia. Aust. A Bat, 34, 649-662.
GREENWOOD, D. R. (1987) Early Tertiary Podocurpaceae:
Megafossils from the Anglesea Locality, Victoria. Aust.
J. Bot. 35, 111-133,
Hitt, R, S. (1982) The Eocene megafossil flora of
Nerriga, New South Wales, Australia. Palaeonto-
graphica Abt B. 181, 14-77,
(1986) Lauraceous leayes from the Eocene of
Nerriga, New South Wales. A/cheringa i(3), 327-352.
LANGr, R, T, (1978a) Some Eocene leaf fragments
comparable to Proteaceae. J. Roy. Soc. WA. 60, 107-114.
162
(1978b) Southern Australian Tertiary Epiphyllous
fungi, modern equivalents in the Australasian region,
and habitat indicator value. Can. J Bot. 56(5), 532-541.
(1980) Evidence of Lid-cells and host-specific
microfungi in the search for Tertiary Eucalyptus. Rev.
Palaeob, Palyno, 29, 29-33,
Martin, H. (1986) On the Philosophy and Methods used
to reconstruct Tertiary Vegetation. Proc. Linn. Soc.
N.S.W. 107(4), 521-533.
Nix, H (1982) Environmental determinants of
biogeography and evolution in Terra Australis. Ch. 5,
In Barker, W. R., Greenslade, P. J. M. (Eds) “Evolution
of the Flora and Fauna of Arid Australia”. (Peacock
Publications, 8, Aust.),
RicHARDS, P. W. (1957) “The Tropical Rainforest”. pp.
xviii, 1-450 (Cambridge Univ. Press, Cambridge).
TRAcEY, J. G, (1982) “The vegetation of the Humid
Tropical Region of North Queensland”. pp, 1-124
(C.S8.LR.O. Melbourne),
Wess, L. J. (1959) A physiognomic classification of
Australian rainforests. J, Ecol, 47, 551-570,
(1968) Environmental relationships of the
structural types of Australian rainforest vegetaion. Ecol.
49, 296-311.
TRZMACRACALVTHUS GEN. NOV. (CESTODA: TRYPANORHYNCHA:
EUTETRARHYNCHIDAE), WITH REDESCRIPTIONS OF
T. AETOBATZDZS (ROBINSON, 1959) COMB. NOV. AND T. BZNUNCUS
(LINTON, 1909) COMB. NOV.
BY IAN BEVERIDGE* & R. A. CAMPBELL}
Summary
Trimacracanthus gen. nov. is erected within the Eutetrarhynchidae Guiart, 1927 for two species,
Prochristianella aetobatis Robinson, 1959 and Rhynchobothrium binuncum Linton, 1909. Both
species are redescribed. The new genus is most similar to Prochristianella Dollfus, 1946 in
possessing a large basal swelling of the tentacle and a distinct basal armature, but differs in having
an asymmetrical swelling and a triad of grossly enlarged hooks on the external surface of the
swelling. The two species are distinguished from one another by number of hooks in each principal
row, size of hooks and number of testes in mature segments.
KEY WORDS: Cestoda, Trypanorhyncha, Eutetrarhynchidae, Trimacracanthus, Prochristianella
aetobatis, Rhynchobothrium binuncum.
TRIMACRACANTHUS GEN. NOY. (CESTODA: TRYPANORHY NCHA:
EUTETRARHYNCHIDAE), WITH REDESCRIPTIONS OF T. AETOBATIDIS
(ROBINSON, 1959) COMB. NOV. AND 7: BINUNCUS (LINTON, 1909) COMB. NOV.
by TAN BEVERIDGE* & R. A. CAMPBELLT
Sunimary
Bevertnor, 1. & CamPBect, R. A, (1987) Trimacracunthus wen, novi (Cestada: ‘Trypanorhynecha:
Eutetrarhynchidae), with redescriptions of 7; aetobaridis (Robinson, 1959) comb. nov, and 7. binuwaicws
(Linton, 1909) comb, ney, Trans, R, Soe 8. Aust, L113), 163-171, 30 November, 1987.
Timacracanthus gen. noy, is erected within the Enutewarhynchidae Guiart, 1927 tor two species,
Prochristianella actobatis Robinson, 1959 and Rhynchobothrium binuncum Linton, 1909. Both species
are redescribed, The new genus is most simitar to Prachristiarielig Dollfus, 1946 1" possessinga large basal
awelling of the leniacle and a distinct basal armature, but differs in having an asyaintetrical swelling and
atriad of grossly enlarged hooks on the external surface of the swelling, The two species are distinguished
from one another by number of hooks in each principal row, size of hooks and number of testes in mature
stgments.
Key Words: Cestoda, Trynanorhyneha, Eutetrarhynchidas, Vrimacracunthus, Prochristiarella
aetobatis, Rhynchodborhrine binuncume.
Introduction
Recent collections of cestodes from South
Australian elasmobranchs included the trypano-
rhynch Prochristianella aetobalis Robinson, 1959
which is apparently a prevalent parasite of some of
the species of rays ovcurting in coastal waters, In
preparing a redescription of the species a number
of significant differences from congeners became
apparent, In addition it was noted that another
species of trypanorhynch, Rhychabothrium
binuncum Linton, 1909, described from a stingray
from the northwestern Atlantic and currently
considered a species incertae sedis (Yamaguti 1959;
Schmidt $986) shared several of the distinctive
features found in P aetnbatis, In this paper we
redescribe both species and show that they consti-
tute a new genous within the Eutetrarhynchidac,
related to Prochristianella.
Materials and Methods
Specimens of P aerobaris were obtained fron the
spiral valves of rays from South Australian coastal
waters, Cestodes were cither washed in sea water,
relaxed bricfly in tap water, then fixed in 10%
buffered formalin, or the spiral valve contents were
fixed with hot formalin and the cestodes were sub-
sequently removed in the laboratory. Cestodes were
* Central Veterinary Laboratories, Sourh Australian
Deparrment uf Agriculture, Institute of Medical and
Veterinary Science, frome Road, Adelaide, 5. Aust. 504K)
| Department of Biclogy, Southeastern Massachusetts
University, North Dartmouth, Massachuserts 2747,
USA,
stained with Celestine blue, dehydrated in ethanol,
cleared in clove oi] ancl mounted in balsam. Six
specimens were also examined by scanning electron
microscopy. Type specimens of PB wetohatis were
borrowed from the National Museum of New
Zealand (NMNZ). All new material collected has
been deposited in the Australian Helminth
Collection (AHC) of the South Australian Muscum,
Adelaide. Additional specimens of FP aétobalis have
been deposited in the British Musem (Natural
History), London (BMNH)}, and the United States
National Museum Helminth Collection, Washington
(USNMHC). Type specimens of R. binuncuen were
borrowed from USNMHC,
Measurements’ are presented in the text in micro-
metres, uiless otherwise stated. The range is
followed by the mean in parentheses. Untess other-
wisé indicated, 10 measurements were made of each
organ,
Terminology for the various elements of try-
panorhynchan-anatomy follows Nollfius (1942) and
Sehntidt (1984).
Trimacracanthus gen, nov.
Generi¢ diagnosis! Eutetrarhynehidae Gulart,
1927. Seolex slender, elongate, acraspedote. Pars
bulbosa and pars vaginalis much longer than pars
bothnidialis. Bulbs long and slender. Tentacle
sheaths sinuous but rot spiral. Prebulbar organs
preset, Retractor muscle attached at posterior
extremity of bulbs, Two bothtidia, margins thick,
free, not contiguous apically, may be noiched
posteriorly suggesting jncipienc subdivision.
Tentacles long with asymmetrical basal swelling
tat |. BEVERIDGE & R.A. CAMPBELI,.
bearing distinctive basal armatiyire, Metabusal
armature heteroacanthous, hetcromorphous,
typidl, Houks solid) Metabasal armature of
allernate half spiral rows beginning with large boak
1 () with recurved point and long base of
implantation, Hooks 2 (29 also large bur smaller
that 7 (')}; remaining hooks diminish in size
becuming slender and shurler as row contimues
towards external face, Strobila acraspedute,
hyperapolytw. Genital pore marginal. Testes
medullary, preovarian, External seminal vesicle
plesen(. Ovary posterior, 4-Johed i) cross section.
Seminal receptacle present. Uterus median. Vitel-
laria cireumcortical
Type species; T qerabutidis (Rabinsan, 1959) comb.
nay,
Other species; 7 Ainurcus (Linton, 1905} comb,
nov,
Trimacracanihus setobalidis (Robinson, 1954)
comb. novi
FIGS 1-15
Prochristianelia aciabarts Robinson, 1959, pp, 49a 394,
figs 26-34,
Types: Holotype from spiral valve of Aerobarus
nurinari Muctler & Henle, 184) (syn. A. renee
caudatus), New Zeukand (precise locality unknown},
March 1955, coll. E. $. Rebinson, holotype NMNZ
ZQO201, paralypes NMNZ ZW 204-206,
Material Examined: from Aetobatus nurinuri: paratypes
NMN4 (ZW 205-6). From /rygonorhina guanerias
Whitley, 1932 (new host record): 23 specimens, Goolwa,
&. Aust, (AHC $2760, USNMEHC 79592), 1 specimen,
Marion Bay, 5. Aust.; 38 specimens, Middleton, S. Aust.
{AHC $2358); | specimen, Port Stanvac, $, Aust. (AHC
52763); 3 speciniens, Cowell, S, Aust; (AHC 52762).
Hrom Mvliohatis austealty Macleay, 1881 (new hivst
rovord); 8 specimens, Middleron, 5, Aust, L[AHC S2455);
| specimen, Young Rocks, Kangarow Island, S$ Aust.
(AHO $2764); Ib specimens, Gerolwa, So Aust. (APC
S27%hy% | specimen, Port Lincaln, § Aust. (AHG S276h).
From Dasyvatis brevicuudatus {Lluttow, 1875) (ew host
revord): LI specimens, Northaven, 3. Ausi. (ALLC $2768)
I specimen, Port Vincent, S$. Aust. (ANC $2769), 1G
Specimens, Streaky ‘Bay, S. Ast. (AHO $2767),
From Dassutiy thetidiy Waite, 1899 (new tiast pecordy +
specimens, Cowell, S. Aust. (ATTIC $2770),
Krom Musrelus antarcsens (Cincher, (R70) (new host
razor): | specimen, Goolwa, S. Agst. (BMNIT 1987.4.7.4).
Desvripien: Slender, delicate worms, up ta 1S nim
Tong, consisting of up 10 |S acraspedote sepments,
Scooles acraspedote (Fig. 2), 2.54-2.99 (2.77) mm
long, 380 530 (410) wade with dao, well scparaled
parelliform bothridia 350-470 (400, n=5) in
diameter. Bothridial margins free, indented poster-
ely, borders thick, lerminaing as short ridges on
internal surface of bothridiym (Fig, 7), Pars
hothridialis 360-460 (420); pars vaginadis 1.2-1.4
(1.31) mm; pars bulbosa 1.18-1.54(1,38) mm, Ralie
of purs bulbosa to pars vaginalis 10095. Tentactes
up to 2.22 mm long, 40-70 (50) tw diameter
excluding hooks; prominent asymmietnical bisa)
swelling, 50-110 (80) wide (Figs. |2-14), Tentacle
sheaths sinuous with some coils, not spiral, Bulbs
at feast eight times longer than wide, 1180-1500
(1300) by 140-180 (160); origin of retractor muscles
ar base of bulb, sucrounded by irregular clumps of
gland cells; prebulbar organs present, Red piemen-
tation anterior to bulbs in living specimens. Pars
postbulbosz very short, 35-110 (67),
Asmature heteroacanthous, heteromorphonus
typical; looks solid. Metabasal armature (Figs 9-11)
consists of alternating, ascending, half spiral rows
of 10-L1 hocks ar mid-tenracle; 12-14 hoaks per half
spiral raw near tip of tentacle. Rows begin on
internal face with hooks | (1') well separated, ending
on eaternal face where hal! spiral rows meet to form
inverted V. Hooks of prinvipal rows begin wilh
single large hook with long base of implantation,
remaining hanks became niore slender with shorier
bases on borhridial and antibothridial faces; hooks
of external face much reduced in size All hooks
become smaller toward tip of tentacle, Hook
dimensions in mid-region of tentacle: 1 (1") arcuate,
stout, 34-46 (43) long, base lengih 20-30 (24);
hooks 2 (2') straighter (Fig, (0), more slender, tip
curved, 38-54 (44) long, base length 36-18 (17);
hooks 3 (3') slender, spiniform with short base
38-50 (46), base 10-18 (16); haoks 4 (4) 38-48 (443
long, base 14-24 (17); hawks 3 (5') 32-46 (40) lang,
base $2-16.415): hooks 6 (6°) 28-40 (32) long, base
10-14 (13); hooks 7 (7'), 18-32 (23) lone; & (89,
12-20 (16) long, 9 (9°)-17' (71'), TH-20 (15) tone.
Basal armialure (Pigs (2-14) consists of eight rows
of hooks on basal swelling of tentacle; hooks
dissimilar on external and internal surfaces. Huuks
of internal surface organized into alternate,
ascending half spiral rows with hooks 1 (1') well
separaled, Proximul mas ene to four consist of
series of large spiniform hooks, gradually
Ueereasing in size as rows continue towards external
surface. Rows five lo cight begin with sloul rase-
thorn shaped hooks | (1); row five continues ins
series of spiniform hooks much stouter than these
of proximal rows one to four. Uall spiral rows one
to five terminate in series of four to cight small
spinilurm hooks on external surflave in inverted
V-formation. Rows six to eight with reduced num-
bers of hooks. On external surface, opposite rows
six to eight, distinct promontory present. sur
mounted by triad of lites cone-shaped hooks with
rounded, festooned bases (Figs 12, 13). Central
hook 24 42 (36) long with recurved point, base
tliameter 18-32 (26), Posterolateral hooks of trind
TRIMACRACANTHUS GEN. NOV. 165
dissimilar, more slender of pair 30-38 (33) long with —_ by 220-404) (300). Genital pores irregularly alternate,
rebated point; base diameter 18-24 (21); stouter in posterior 1/3 of mature segment, 67% of
hook 30-36 (33) long, base 16-24 (21). segment length from anterior end; 75% of segment
Mature segments 1.19~3.09 (1,99) mm by 180-310 = length from anterior end of gravid segments.
(260); detached gravid segments 2.30-4.21 (3.11) mm = Genital alrium small. Cirrus sac globose, 70-140
Figs 1-7. Trimacracanthus aetobatidis (Robinson). 1, entire cestode; 2, scolex; 3, mature segment; 4, female genitalia;
45, gravid segment; 6, cirrus sac with external seminal vesicle; 7, bothridia, Scale lines: fig. 1, 8, 1.0 mmy figs 2-4,
6-7, OL mm.
166 1, BEVERIDGE & R. A. CAMPBELL
ee
z ip ma
a nV i, re MS)
vy a 4 jf ip C if y x
oe Vif fs
f i
es
1 ff ey GE IK
A Ww fy See. betes las
oh ( (/ Pass, wa js e . pai i
in
(on
et
Figs 8-14. Trimucracanthus uctobatidis (Robinson), leniacular armature. 8, bothridial surface of tentacle, 0,1 nm
from tip of fully extended tentaele; 9, bothridial surface, metabasal region, 0.8 11 from base; 10, antibothridial
surface, metabasal region, 0.8 mm from base; 11, external surface, metabasal region, 1.9 mm Irom base; 12, basal
neaion of tentacle, bothridial surface; 13, basal region, external surface; 14, basal region, inlernal surface. Seale
ine: 0.1 mini,
TRIMACRACANTHUS GEN, NOV. Is?
(100) by 3ie-t1O (8D); civrus unanmed. External
seminal yesicle pyriform (Fig. 6), jdnate te cirrus
sac; vas deferens coiled, located in midline between
cirtus sac and ovary. Testes preovarian, sub-
spherical, 50-100 by 40-50, forming Iwo parallel
renwss testes tLumber 68-85 (76) per segment; testis
distribution: postvagingl and poral 6-8 (7)
prevaginal and poral 28-35-(31); antiporal raw 34-43
(38), Vagina passes anteroventral to cirrus pouch,
turns sharply posterior in midvedtral line and
descends fo ovary, Vagina divided into Ovo
segments; first scement greatly expanded from
penital atrium (6 anterior level of ovary Where Sperm
are stored, wall lined by simple epithelium; con.
stricts forming second segment, narrow sperm duct
passes Ventral td ovarian isthmus and joins ootype.
Ovary bilobed in dorsoventral view, lobes suberual,
130-270 (190) by 30-60 (40); Mehlis' gland 80-220
{i50)in diameter, Vitelline follicles cireumeortival,
BO in diameter. Uterus sacciform in detached gravid
segments, filling entire preavarian medullary region,
Trimacracaniines binuncus (Linton, 19049)
comb. nov.
FIGS 16-215
Rhyachohotinien binuacum Linton, 1909, pp. 181-182,
fins 5$-f4,
Types! Lectatype from spiral valve of Dasyatis say
(Le Sueur, 1886), Tortugas, Florida, USNMHC
No, 9010; paraleciolypes, USNM SOIC,
yen examined) trom Dasyaris say; 4iypes, USNMHC
on
Deseription: Small, delicate warms, 4.9-6.4
é7i mm (n=2) long, consisting of 5-6 (n=4)
actuspedote scements, last segment comprising 65%
er more af Lotal length of worn (Fig 16). Scolex
acraspedote (Fig. 15), 1.30-1.65 (1.53) mim (n=4)
long by 190-280 (248) (n=3) wide, with two
rounded, Well separated, bothridia, 175-250 (216)
{n-4) long by 160-250 (203) (n-3) in diameter.
Buthricial margins free, borders thick, no indenta-
tion or incipient division on posterior barder. Pars
hothridialis 175-250 (223) (n=4); pars vaginalis
800-960 (853) (n=4); pars bulbosa 750-830 (787)
(n= 4), Ratio of pars bulbosa to pars vaginalis (1.09.
Tentacles extended ta 550 bur nor fully everted.
Tentacle widths. excluding hooks, 80 wide al basal
swelling in lareral view, lower metabasal diameter
§5, mid-tentacle diameter 45, tp diameter 45.
Tentacle sheaths sinuous with few coils. Rulbs
clongate, 770-830 (778) (n=4) by 70-108) (83) (n=4},
Tetractor muscle originales at base of bulb,
surrounded by irregular mass of gland cells;
prebulbar organ present. Pars postbulbosa absene.
Armature heteroacanthous, heteromornhyis,
lypical; hooks solid. Metabasal armature (Figs 19,
20, 22, 23) consists of alternating, ascending, halt
spiral rows of 12-14 hooks each, Rows begin on
internal surface with haoks } (1) well separated,
ending on external surface m inverted V-formations,
Hooks | (l'} Jarge. falciform, with extended
posterior lieel; remaining hooks of principal cows
straighter, falciform then spiniform, eradually
decreasing in size. Hook dimensions in mid-region
of tentacle: 1 (1') stout, stronely recurved, with
extended beel, [6-32 long, base length 15-20, heel
4-5 long, height 9-23; hooks 2 (2') similar 10 1 (Lt)
except. straighter and with reduced hase, length
19-20, base length 1-12, height 10-\1; hooks 33")
spiniform, length 15-18, base length 6-8, height
10-11; houks-4 (4')-9 (9') decreasing in length from
25-2 hooks 10 (10')-14 (14°) only 17-18 long.
Basal arinature (Figs 21-23) consists of six rows
of hooks on hasal swelling of tentacle; hooks dis-
Similar on external arid internal faces. Hooks of
internal face organised into aletnate ascending half
spiral rows with hooks | (h') widely separated. Rows
hegin with stout falciform hooks with long postesor
heel, onented transversely; remaining hooks of
proximal rows spiniform, eradually decreasing in
Mize as rows continue toward carerna!l surface,
Smaller spiniform hooks on external face meet in
inverted V formations (Fig. 22). Distinct promon-
tory on external face surmounted hy triad of large
cone-shaped hooks with rounded, festagned bases
and técurved points (Fig. 18), Foremost hook of
triad largest, length 34-42, hase diameter 18-25,
Posterolateral hooks of triad unequal; hook
adjacent buthridlal side larger, length 28-38, base
dhameter 16-218; hook adjacent lo antibothridial
face smallest of triad, length 24-28, base diameter
$-12.
Malure segments 1.497-1.27 (L.L7 mm) (n=3) long
by 150-210 (187) (n =3) wide; tetminal seemetits,
3,12-4,45 (3.67 mov (n=4) long by 200-400 (300)
(n=4) wide, nangravid. Genital pores irregularly
ullernate, in posterior half of mature segment,
Detached eravid segments not present. Cierus sac
pyriform, 60-90 (69) (n=4) by 23-34 (29) (n=4),
external seminal vesicle 40-50 {n—2) in diameter,
adnate to cirrus sac. Vas deferens coiled, forming
loop, extending in mecial line postenorly to level
of ovarics. Testes preovarian, ovoid, dimensions
60-70 wide by 20 long, forming owe parallel rows,
Testes number 105-109 (107) (n=2) per segment,
testis distribution, postvaginal, poral 13-15 (n= 2);
prevaginal, poral 35-39 (n=2), antiporal 54-56
(n=2). Ovary bilobed in dorso-ventral view; lobes
stibequal. 240-260 (n=2) by 160-220 in=2). Vagina
forms seminal receptacle #0 in diameter, Mchilis'
gland well developed, pastavarian, Vitelline follicles
circumeorical, continuous frau anterior ro
posterior margins of segment.
168 1, BEVERIDGE & R, A. CAMPBELL.
Figs 15-17. Trimacracanthus binuncus (Linton). (5, scolex; 16, terminal segment; 17, entire worm, Scale lines: fig.
17, 10 mm; figs 15-16, 0.1mm.
169
TRIMACRACANTHUS GEN, NOY.
-23. Trimacracanthus binuneus
19, métabasa
surface; 22,
external surface of basal swelling;
#; 20, metubasal region, internal surface to left; 21, basal armature, bothridial
surface; 23, basal and metabasal armature, internal surface;
(Linton). 18, triad of enlarged hooks on
| region, antibothridial surfac
basal and metabasal armature, external
note long guards on hooks 1 (1') of metabasal region. Scale line: 0.1 mm,
igs (8
(70 |, BEVERIDGE & R. A: CAMPHELI,
Discussion
The 1wo species described above possess a combi-
uation of characters pot found together in other
euletrachyichid cestodes, in particular, an
asymmetrical basal swelling ataied with a triad of
laree hooks und « metabasal armature composed
of ascending hal{-spiral rows of hooks with hooks
which diminish in sive from the internal to the
exterial surface suggesting that a new genus is
needed to accommodate them.
Dollfus (1942) assigned twu genera lo the Euletra-
rhynchidae. namely Evfetrardpynchus Pintner, 1913,
and Christianella Giuiart, 1933. He Jater (1946)
added Diesingefla Gutart, 1931, and Terrerhyncho-
borhrium Diesing, 1850 and created two new génera,
Prochristionetla Dollfus, 1946 and Purachristianella
Dollfus, (946 bringing the total number af eutetra-
rhynchid genera to six. Dollfus (1969) subsequently
removed Tetrarhynchahothriaen from rhe
Eutetrarhynchidae and made ir the type genus of
anew family, Tetrarhynchobothridae Dollfus, 1969
which remains monotypic, Mecistoholhriue Heinz,
& Dailey, 1974 is a more recent addition to rhe
family Eutetrarhynchidac. Unfortunately Diesin-
gellaand Christionedia remain very poorly defined
and require s thorough redescription to be properly
included in current systematics. Mecistoberhriian
Appears, from its description (Heinz & Dailey, (974),
to be very similar to Renibulbus Feigenbaum, 1975,
originally placed in its own family, Renibulbidae
(Feigenbaum, 1975). Subsequently, Renibulbus was
placed as a synonym.of EulerrwrAyncus along with
Tetrarkynchohatheitin by Schmich (1986). Schmidt
{t986)} also added Oncomegas Dollfus, 1929 to the
family,
Linuls of the genem, Eurerrarbyachyus, Prociris-
tenella and Parachrtstianelle are not clear and this
has resulted in confusion in the allocation of species
to genera, In defining the genus Prochristianelle,
Dollfus (1944) emphasized (wa principal huravters;
the inuredse and subsequent decrease in metabasal
hook size from internal to external surfaces in Pro-
eheistranella conipared with a devline th hook size
in Pucvetristianetiaz ind the presence of a tentacular
basal swelling with a characterisae armature in
Prochristianelia hut not in Parachristianelia, He
placed two species in Prochrisrianetia, the type
Species, feveanicaia with a heteromarphays
merabasal armature and A tenvispine (Linton, 1890)
with a homeomorphous armature. Dollfus (1946)
defined Purachristianella as having a decrease in
hook size from internal co external surface and
jacking a basal swelling of the tentacle. Sub-
sequently, Kruse (1959) added twe new species to
the genus which conformed closely with Dolifus’
definition, Yamacutl (1959), however, lgaored the
nature ol the metabasal armature In his key to
genera (of the fumily relying instead On the presence
af a basal swelling afd it is evident from the
lireralure jhal some workers have followed
Yamaguti’s key rather than Dollfus’ orgimal
diagnesis. OF specific concern here is Robinson’s
(1959) description of Provhristianella actobaris from
New Zewland which combines the metabasal
armature of Patachrisnanelfa with the basal
arinalure on a tenfacular basal swelling like
Prachrisnareila, Following the key of Yamaguti
(1959), Robinson (1959} placed the species in
Prachrisiivenella bur with some Nesitatiun, as
Museum records show he had originally considered
use 4 ew generic name for the species, Thenacna-
canthus, We consider that the bwo species desorbed
above fir neither Prachristianelfa nor Perachristia-
nella, and that a new genus is required to accom-
modate them, for which we propose f6 use (he name
initially Suggested by Robinson in notes attached
to ihe jype specimens,
Distinctive characters of the new genus Trimacna-
centhas are the combination of a mbetabasal
armature similar ta Parvchristiamella but with a
prominent, asymmetrical tentacular basal swelling
bearing @ distinctive armature including rhe
presence of a triad of enlarged hooks. The two
species placed in the new gtnus are T, aelabatidis
and 7 binvacus, T. binuacus was originally
described by Linton (1909) and regarded by
Yamnagurl (1959) and Schmidr (L986) as incertie
sedis. The syntypes of RAynchobothruam dinuncum
(USNM 9010) were examined and the oncotaxy and
analomical leatures were found to be very similar
to but distinct from those of 7 aetobatidis. Both
species possess a metabasal armature al the typical
heteroacanthous type in which the haok sizes begin
with very large hooks then decrease in size towards
the external surface. Also, both species have a
pronounced asymmetrical basal swelling with a
distinct basal armature. There are three cone-shaped
hooks on. the external fave of the basal armature
in both species, 7 gerebatidis may be distinguished
from 7 dinuiicks by the following features: number
of hooks per principal rove at mid-lennacte (10-11
versus 12-14); hooks £ {1') in the principal rows lack
an extended posterior heel in T wetidaetidiy bul a
heel is pronounced in TL Bbinyncus; 2 sharp
transition In hook size on che external lace (7
acinhatidis) versus a gradual decrease in hank size
17. binuneys); testis number per sezment (68-85 in
T. aeioberis versus 105-109 in To bintuncus); number
of segments (1S versus 6); nonspinaus scoles and
strobila of 7 aerobaridis versus aspinous condition
in T. Ginweteus, and presence of a short pars
posthulbosa in 7 eelobetidis which 7 dvnaneus
lacks.
TRIMACRACANTHUS GEN. NOVY. 171
Robinson (1959) used two names, P gerobatis and
P. aetobatidis, in his description of the species. As
the host genus is Aefobatus, we assume that P
aetobatis is a typographical error, and have adopted
the name aetobatidis.
Southwell (1929) reported a single specimen of
R, binuncum (as Tentacularia binunca) from
Dasybatus? walga from the “Ceylon Pearl Banks”,
and. reproduced Linton’s (1909) figures of the
species. The whereabouts of Southwell’s specimen
is not known, and hence its identity cannat be
ascertained.
Acknowledgments
Thanks are due to B. G. Robertson, R. R. Martin
and M. G. O'Callaghan for collecting specimens
and to J. R. Lichtenfels (USNMHC) and R. L.
Palma (NMN7Z) for the loan of type specimens.
This study was supported financially by the
Australian Biological Resources Study.
References
DoLiLFus, R. (1942) Etudes critiques sur les
Tétrarhynques du Muséum de Paris, Arch. Mus. nat.
fist. nat,, Paris 19, 1-466.
(1946) Notes diverses sur les Tétrarhynques. Mém.
Mus. nat. Hist. nat, Paris 22, 179-220.
(1969) De quelques cestodes tétrarhynques (hétéra-
canthes et pécilacanthes) recoltés chez des poissons de
Ja Méditerranée, Vie et Milieu, Sér, A 20, 441-514.
FEIGENBAUM, D. L. (1975) Parasites of the commercial
shrimp Penaeus vannamei Bourne and Penaeus brasil-
fensis Latreille. Bull. Mar. Sci, 25, 491-514,
Heinz, M. L. & DaiLey, M. D. (1974) The Trypano-
thyncha (Cestoda) of elasmobranch fishes from
southern California and Northern Mexico. Prac.
Helminthol. Soc, Wash. 41, 161-169.
KRUSE, D. N, (1959) Parasites of the commercial shnmps,
Penaeus aztecus Tves, P. duerarum Burkenroad and
P setiferus (Linnaeus). Tulane Stud. Zool. 7, 123-144,
LINTON, E. (1909) Helminth fauna of the Dry Tortugas.
I. Cestodes. Carnegie Institution of Washington
Publications 102, 157-190.
ROBINSON, F, S. (1959) Some new cestodes from New
Zealand marine fishes. Trans, R, Sec. N.Z, 86, 381-391.
Scumint, G. D, (1986) Handbook of Tapeworm Identifi-
cation. (C/R.C, Press, Boca Raton, Florida.)
SoOuTHWELL, T. (1929) A monograph of cestodes of the
order Trypanorhyncha from Ceylon and India, Part I,
Spolia Zeyland. 15, 169-312.
Yamacuti, S, (1959) Systeme Helminthum, The
Ci ay of Vertebrates, Vol. 2, (Wiley Interscience, New
York.
j
i GER
chee me uae a! is
TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
VOL. 111, PART 4
CHANGES IN SALINITY AND IN THE DISTRIBUTION OF
MACROPHYTES, MACROBENTHOS AND FISH IN THE COORONG
LAGOONS, SOUTH AUSTRALIA, FOLLOWING A PERIOD OF
RIVER MURRAY FLOW
BY M. C. GEDDES*
Summary
This study was carried out to investigate the effects of the above average flow in the River Murray
in 1983-84 on the salinity and the distribution of the biota in the Coorong Lagoons. Salinities fell
only slightly in 1983, but by mid 1984 salinities had fallen significantly to be brackish (< 30°/oo) in
the North Lagoon and moderately hypersaline (55-70°/oo) in the South Lagoon. The biology and
distribution of the biota showed changes that correlated with the freshening of the Coorong. Ruppia
megacarpa flowered profusely in the North Lagoon in October 1983, and R. tuberosa grew
extensively in the South Lagoon in 1984. The “estuarine-lagoonal” macroinvertebrate fauna in the
North Lagoon extended its distribution to the southern extent of that lagoon; the species richness
remained low with only two previously unrecorded species being collected. None of the typically
estuarine-lagoonal macroinvertebrates occurred in the South Lagoon within the study period,
although salinities were generally within their tolerance range in winter-spring 1984. Some
freshwater fish occurred in the North Lagoon and Aldrichetta forsteri and Acanthopagrus butcheri
(estuarine-marine species) moved into the South Lagoon for a brief period in spring 1984.
KEY WORDS: Coorong Lagoons, salinity, macroinvertebrates, fish, Ruppia.
CHANGES LN SALINITY AND IN THE DISTRIBUTION OF MACROPHYTES,
MACROBENTHOS AND FISH IN THE COORONG LAGOONS, SOUTH AUSTRALIA,
FOLLOWING A PERIOD OF RIVER MURRAY FLOW
by M, ©, Genaes*
Summary
Geopes. M. C. (1987) Changes in salinity and inthe distribution of macrophytes, macrobenthns and fish
in the Coorong Lawoons, South Australia, following u period of River Murray flow, Trans, R, Sov.
S. Aust, 114), 173-181, 30 November, 1987.
This study as carried out td investigate the effects of the above average flow in the River Murray
in 1983-84 on the salinity and the distribution of the biota in the Coorong Lagoons, Salimbhes fell anly
slightly in 1983, but by mid 1984 salinities had falleu siznifivantly to be brackish (<30%o0) ili tlie Nortli
Lagoon and moderately hypersaline ($59-70%ov) in the South Lagoon. The biology and distribubon af the
biota showed changes that correlated with Lhe freshening of the Coorong. Ruppia megacerpa Flowered
profusely in the North Lagoon in October 1983, and R, tuberosa grew extensively in the South Lagoon
in 1984. The “estuarine-lagoonal” macroinvertebrate fauna in the North Lagoon extetied its distribution
(o the southern extent of that lagoon; the species richness remained low with only two previously unrecorded
Species being collected. None of the typically estuarine-lagoonal macroinvertebrates ogcurred in the South
Lagoon within the study penod, although salinities were venerally within their tolerance range.in wintet-spring
1984, Some freshwater fish occurred in the North Lagoon and Aldrichetta forsteri and Acanthopagrus
butcheri (estuarine-marine species) moved into the South Lagoon for a brief period in spring 1984.
Key Worvs: Coorong Lagoons, salinity, macroinvertebrates, fish, Ruppia.
introduction
The Coorong is an elongate coastal lagoon
systeot which extends from the mouth of (he River
Murray some I) km southeast along the South
Australian coast (Fig. 1) and is characterized by a
preal rarige of salinity Muctuations (Geddes & Butler
1984). Exchange of water into the Coorong is from
ihe Murray Mouth region at the northern end of
the Coorong. Here eilher sea water from Encounter
Bay or fresh water from the River Murray via Lake
Alexandrina, can enter the Coorong. ‘The formerly
estuarine nature of the Murray Mouth region has
heen changed by River Murray regulation and by
the erection of (the River Murray Barrages, a system
at low levees and gates (approximately 600 in all)
actoss the outflow from Lake Alexandrina (Fig. 1),
Ar times of low River Murray flow the water
entering South Australia is sufficient only to meet
abstraction requirements and evaporative losses and.
al these times the gates of (he barrages are shut and
the barrages separate fresh water retained in Lake
Alexandrina from sea water in the Murray Mouth
gestion. AL periods of moderate or high River
Murray flow, the gates on the barrages are opened
to varying degrees and fresh water flows into the
Murray Mouth region and the Coorong. Therefore
* Depar(ment Oo) Zooloey, University OF Adelaide, GPO,
Box 498, Adefuide, S. Aust. S00).
the flow conditions in the River Murray determine
the salinity of the water in the Murray Mouth region
which is available for mixing into the Coorong and
thus are a major factor controlling salinity in the
Coorong Lagoons. The nature and distribution of
the biola of the Coorong Lagoons is influenced by
salinity (Geddes & Butler 1984; Kangas & Cieddes
(984) and so River Murray flow is likely to influence
ihe blology of the Coorong Lagoons.
A previous study of the Coorong (Geddes &
Butler 1984; Kangas & Geddes 1984) was. under-
taken during u d6-month period (Decenther 1981
to March 1983) of no ourtlow trom the barrages.
During this time the Coorong's salinities were
marine to moderately hypermarine in the North
Lagoon (35-50°%no) and strongly hypersaline
(80-110°/o0) In the South Lagoon, These conditions
greatly restricted the distribution of the marine-
derived estuarine-lagoonal fauna in the Cocrung.
(n 1983 and 1984 there were moderate to high flows
in (he River Murray; the Murray flow year }983-34
recorded a flaw of 8,08 million MI at Lock 1
compared with the long term median (1949-50 to
1982-83) of 3.96 million MI, In mid 1983 there
began» considerable ont low trom the barrages 50
that the Coorong received diluting flows. This paper
describes the changes in the salinity patterns in the
Coorene fram March 1983 until March 1985 snd
the effects of these changes on the distributions of
muctuinvertebrates, fish and macrophytes in the
North and Sourh Lagoons.
174
M. C. GEDDES
Monthly
Borrage Flow
8 A 20930 40 5a 6O ef 10 780
c
5m og hot sampled 12-12-83
Mureay Mouth gy
(Mix lav§}
Are North Sugeon South Lagoon
wsdertdetbeed lepedosredberidscrederpehmarsionrcinchst = = = ateeceeeneenneeenenecvacseuesiesenies
7-3-83
TTT Ti
$ 20 a fao 50 lac fo 80
77-84—
15m 26°7'84
Buy
io fs fa 30 5a 55 eo Jas 790
25 As 20:9: 84-
18-10-84
ur Sour
area Nuria Voguun South Lagaan
1983 1984 1985
COORONG BIOLOGY AND RIVER MURRAY FLOW 175
Methods
Eight visits, at approximately three monthly
intervals, were made to the North and South
Lagoons from October 983—March 1985. The visits
to the two lagoons were generally within twa weeks
of each other. In the North Lagoon physicochemical
measurements and bivlogical samples were taken
from (he nine stations of Geddes & Butler (1984).
The methods for measuring conductivity at 25°C
(Kos), salinity (measured as tocal dissolved solids,
TDS), chlorophyll, turbidity, Secchi dise trans
pareney for Secchi depth) and OQ) concentration
and for collecting benthic and littoral invertebrates,
fish and macrophytes were as in Geddes & Butler
(1984). In the South Lagoon K 5, TDS and
{urbidities were meusured and benthic and littoral
invertebrates were collected from seven stations
(Fig. 1) The first of these stations (1S) was al
Magrath Flal which is within the constricted region
herween the North and South Lagoons. The other
six were in the South Lagoon from Vila de Yumpa
to just north of Tea Tree Point. Additional
information on the distribudien of chironomids,
macrophytes and fish in the South Lagoon were
provided by Dr David Paton (Zoology Department,
Umyersity of Adelaide) and Mr David Hail
(Department of Fisheries, Sourh Australia). Data
on estimaled olflows from the River Murray
Barrages and on the patterns of opening and closing
of the gates ol’ the hatrages were oblained from the
Engineering and Water Supply Department,
S. Aust,
Results
Physicochemical
Longitudinal and vertical patterns in salinity
(TDS) inthe North and South Lagoons, estimated
outflows from the River Murray Barrages to the
Murtay Mouth and Coorong, and pitlerns of
opening and closing of the barrages during 1983
and until mid 1985 are showitin Fig. 1. The barrages
were closed fram December 198] (see Geddes &
Butler 1984) until f July 1983. In March 1983
salinities Were hypermanne throughout the Coorong
ranging from 40°/%o0-130°/o0 (Fig, 1A), By October
1983 (Fig. 1B), after 3 months of barrage outflow
around | million MI per month, salinities had fallen
and ranged from 25 to 60°/00 in the North Lagoon
and Were about 70°/oo in the South Lagoon. By
December (Fig. 1’), alter cominued good Maws,
salinities at the northern end af the North Lagoon
had fallen slightly while those at the southern end
of the North Lagotn had tisen, There were
considerable vertical salinity gradicnis at the
northern and southern ends of the North Lagoon
presumably caused by deeper level incursion of sea
water and highly saline South Lagoon water respec-
tively. A major freshetiing of the North Lagoon
occurred helween December 1983-and March |984
{Fig. 1D) with the entire lagoon becoming brackish
and the northern half having surface salinities below
S*/ou. There were sharp vertical salinity pracients
as fresher water overlaid denser saline water.
Ourflows over this period were moderale (approx.
0.3-0.8 million MI per month) and by March the
barrage gates were mosrly closed. Iu the South
Lagoon there was a longitudinal salinity gradiene
from 90+|40°%oo presumably as a result of high
evaporation at the shallow southern extremity and
limited tongiludinal mixing, By May-June (Fig, 1B),
after a continued period of moderate flow (0.3-0.4
million MJ), an evea longitudinal gradient. from
<5-25'7o0 had developed in the North Lagoon
and there was a steep gradietit between the two
lagoons, although there had been some freshening
of the South Lagoon. In June and July mauwthly
oulflows were low and sea waler moved from the
Murray Mouth inte the North Lagoon, Further-
More, by July 26 there had been a consilerable
movement of saline water from the South Lagoon
into the North Lagoon (Fig_ 1¥), presumably as a
result of changes in the water levels in the lagoons.
This is clearly shown in Fig. 2, where lofigitudinal
salinity patterns show that salinities rose from
24%o0 and 41°%no at Stations 9 and 1S on June 5
to 24°/op and 72° on June 14, This exchange
between the lagoons broke down the steep salinity
discontinuity thit had existed between them, During
August and September of 1984 (Fig. 1G) barrage
Outflow increased and the North Lagoon shawed
an even longitudinal salinity gradient from
<10-3)%oo, Exchange from the North Lagoon
and seasonal rainfall further freshened the South
Laguon. High outflows (>15 million MIL per
month) were sceorded in Oeteber and early
November, but by carly December all gates on the
barrages were closed, This reduced flow allowed
seawater to enter the Coorong so that on December
4 (Fiz. 1H) the longitudinal salinity pattern in the
Norih Lagoon showed a block of fresher (<10%o0}
Ti. tL. Longitudinal and vertical patterns in salinity (DS oo) in the North and South Lagoons of the Coorong
al ¥ sampling times from March 14983 wntil March [985 and estimated monthly barrage outflow from the River
Murray, The sampling localities ia the North Lagoon (1-9), in the constricted region between the lagoons {1S},
und in the Sourh Lagoemn (25-75) are indicated, Closure of the burrages is indicated by the solid harizonral bats
in the Murray Mauth region, The degree of opening of the main barrages ar Goolwa (Gi) and Tauwiteherie (1)
ts Indicated hy the pesirion and size of the breaks in the bars representing the barrages.
196
(a
Mes}
fas
ab
O
i
any
1P 4 45STH OIE
Srotion
223545 $55
Fig, 2. Longitudinal patterns of surface salinity on Sand
{4 Jume 1984. Note the sharp gradient between Station 9
and Station 25 and the movement of saline water into
the région 28 and 9 on June 14.
wafer half way down the lagoon bounded by saltier
water ta the northern and southern cnds, All
barrages remained closed until the final sampling
date in March 1985 (Fig, 1§) by which time salinities
in the North Lagoon had risen above 35°%on and
thase in the South Lagoon to 140°/o0; so that the
salinity pattern for the Coosong was very similar
to that in March 1983.
Water temperature in. the North Lagoon showed
a seasonal pattern front a high of 24°C to a low
of 11°C, On calm days a small vertical gradient
existed, with surface temperatures up to 15°C
higher than bottom temperatures, Secchi dise trans-
parencies were high (up 10 2.2 m) at the northerly
stations in Lhe period of no flow in March 1983 and
then fell co around 0.5 m in October. Subsequently,
they fluctuated generally between 0.5 and 1.5 m with
fo apparens pattern, probably reflecting, local wind-
driven turbulence. In the South Lagoon Secchi disc
Iransparencies were lower varying between 0.3 and
6.7 m
Biological
Chlorophyll a levels in the North Lagoon showed
a longitudinal increase from Station 1 to 9 in March
1983. Diatoms, along with flagellates at Statyons 7
and 9, were the dominant phytoplankters, During
M. ©. GEDDES
the flow period from mid 1983 until late 1984
chlorophy! levels fluctuated usually between 5 and
30 mg m7? but higher levels were recorded at
Stations | and 3 when Planktonema lautabornei
was the dominant phytoplankter (presumably a
wash-out from Lake Alexandrina where it is usually
abundant (Geddes 1984)}). By March 1985 chloro-
phyll levels were tow al Stations 1 and 3, and high
at Stations 7 and 9 where Mazellates bloomed.
During 1983 and 1984 Ruppia megacarpa
continued to be the daminan{ macrophyic in the
North Lagoon with Lepilaena cylindricurpa also
common and Zostera muelleri common at. the niore
northerly stations. The Ruppia and Lepilaeng beds
were vigorous, extensive and Mowering profusely tn
October 1983 along the length of the North Lagoon,
especially from ‘Station 5 southwards, They
remained extensive until June 1984 and then died
back. They became extensive and vigorous again
by December 1984 hut no flowering was observed
in October or December 1984,
In the South Lagoon Ruppia tuberosa was the
dominant macrophyte with Lamprothaninian
sometimes common, Over the period Octeber 1983
until July 1984 small specimens of Ruppia were
observed in restricted areas, By September 1984
extensive beds of Ruppia occurred throughout the
South Lagoon, These were probably produced from
seeds thai germinated in the freshening water
(D, Paton pers, comm,), In December the northerti
and middle areas of the South Lagoon were fringed
with extensive beds af Ruppia growing thickly in
the shallows and flowering profusely. Ruppia died
back during summer as water Jevels dropped and
salinity rose. By April 1985 Ruppia was recorded
only in the northern half of the Lagoon to Station
6S.
The macroinvertebrates and fish collected in the
North Lagoon are listed in Table |, Most species
were collected on most occasions at Stations | ty
5 where salinities usually ranged from 5-45°/o0,
although some species including the polychactc
Australonereis ehlersi, the gastropod Salinalar
fragilis and the bivalve Nefospisula trigonella, were
much less common (han in the earlier study (Geddes
& Butler 1984), In March 1984, when salinities fell
to 2°%%oo-al Stations 1.and 3 and 5°/oo at Station 5,
the polychaetes Cergtonereis aequisetis (formerly
C. pseudoerythraensis), Nephyis australiensis and
Capitella capitita were not collected from Station |
although C. dequisetis was present at Station 3 and
all three species were present at Station 5. Numerical
dominant species thal remaincd in high abundance
in che littoral area during the estuarine phase in the
North Lagoon were the amphipods Melita zey-
fanica, Paracorophiim and Megamphopus, the
COORONG BIOLOGY AND RIVER MURRAY FLOW iW
TABLE 1. Mucroinvertelrates and fish collected in hand net and benthic grab sumples
tram the Nortit Lagoon of the Coorong from October 1983 until March 1985, (Those
marked * were not recorded when similar samples were taken in 1982 (Geddes & Butler
1984),)
Fish
Arenigobius bifrenatus (Kner)
Pseudogohius alorum (Sauvage)
Atherinosoma microstoma (Ginther)
* Philypnodon grandiceps (Krefft)
* Pseudaphriris urvilti (Valenciennes)
* Nematalosa erebi (Giinther)
Decapods
Amphipods.
(Crustaceans
Macrobrachium intermedium (Sumpsan}
Melita zevlanica Stebbing
Paracorophium cf. excavatum
' Megamphopus/Podaceropsis/Gammaropsis
Ostracod
Polychaetes
Osticythere reticulata Hartmann
? Ceratonereis aequisetiy (Augener)
Nephiys australiensis (Fauchald)
Australonereis ehlersi (Augener)
* Privnospio cirrifera (Wiren)
Ficopomiatis enigmaticus (Fauvel)
Boccardia chilensis Blake & Woodwick
Cepitella capitata (Fabricius)
Capiiellides spp.
Fabriciinae
* Questidae
Gastropods
Hydrobia buccinoides (Quoy & Gaiman}
Salmator fragilis (Lamatck)
* Tafea rufilabris (Adams)
Bivalyes
Notospisula trigonella (Lamarck)
Arthritica semen (Menke)
Soletellina donaciaides. Reeve
Chironomid
Eplhiydrid
Insects
Tanytarsus barbitarsis Freeman
Ephydrella sp-
' This population was identified as Megamphapus sp. in Geddes & Butler (1984) and
Kangas & Geddes (1984), but is more properly assigned to this multi-genus group,
which is in need of tevision (J. L. Barnard pers, comm.). Megamphopus sp. 15 used
elsewhere in this paper for brevity.
? This population was identified as C. pseudoerythraensis Hutchings in Geddes &
Butler (1984), but has been synonymized with C. aeguisetis (Augener) (Hutchings &
Glassby 1985).
polychaetes Cerutonéréis weguisetis and Fica-
Ppomatus enigmaticus, and the gastropod Hydrobia
buccinoides. The daminants in the benthic samples
were Capitella capitata and Paracoraphium with the
bivalves Notospisula trigoneHa and Arthritica
semen and the polychaetes Nephtys australiensis
and Prionaspio cirrifera common,
At Stations 7 and 9 high salinities (54-740) in
March 1983 restricted the fauna so that only
Cuapitella captieta, dipterans and the hardyhead
Atherinosoma microstome occurred at Station 9,
and thesé plus Sulinator fragilis and Hydrobia
buccinoides at Station 7 (Fig, 3). By December 1983
after salinities at Station 7 had fallen below 50°co
in October, Paracorophium, Megamphopus and
Capitellides were present and this last was also
collected from Station 9, When salinities dropped
sharply in March 1984 Ficopomatus enigmaticus,
the fabriciine polychaetes, Arthritica semen and
Pseudogobius olorum were collected from
Station 7, and then Melita zeplanica, Ceratonereis
aequisetis and Notospisula trigonella appeared in
June and Prionospio cirrifera, Macrabrachiunm
Intermedium and Tateq rufilabris appeared Jater.
Thus by October 1984 almost all of the common
estuarine-lagoonal species that occur in the
Coorong had colonized Station 7. The same pattern
occutred at Station 9 except that colonization was
generally later and Melita zeylanica, Tatea rufilabris
and Macrobrachium intermedium were not found.
When salinities rose from December 1984 to Mareh
1985 many species disappeared from Stations 7
and 9 (Fig. 3).
None of the common esluarine-lagoonal inverte-
brates of the North Lagoon were collected from the
South Lagoon. Here, the macroinvertebrate fauna
was very restricted, comprising only the chironomid
Taawtarsus barbitarsis, the ephydrid Ephydrella,
other dipteran larvae, the isopod Hal/oniscus, searler
and ostracods including Diacypris compacta.
ITS M,C, GEDDES
Metin sevlorice
Paracoraphiam Vt arcane
Mecomnphopus
Caraaner eis ceqoiiel!:
—_— Pranespia cirnterc
Copitelia woptdta
Ficopomars eatgmoticus
Copiteiides sop
Fodric now
bydrabie Aucoinedes
_ Tateg nistiigbris
hoa Sain ator tragiis
Notasoiswa Ieonelia
Arthihed semen
———en._ Athorinzame mecrastama
— Prevdegdéivs alorum
+ Mecrebrochram intermedium
— Jonytorsus hartitarsis
—— Fphyareila sp.
ADOT TOM
rai
Bs (hu,
Vr
v NFOLRT;ASANGIERANSUASORDUTWA
{253 1954
_ Morecorophwin of sacwvorum
S-arjon J Megomphopus — ;
= Cerchangrins néguisens
_ $name HIPPO.
Pripaasgia currtsee
Fxopdmatus esrigmarious
Comjelhdes spp
Fabriaiince
Hyorobia burcinaides
+ Sgnnatee fragilis
Nerespisiia tigonetict
Artartica semen
Atherinosoma imicrostana
Pseudegohius alorum
+ — Tanytorsus borbitarsss
+ Eptiydreiia sp.
BOTTOM
TUF
+
4
Od er ert
SEMAN ISM SONTIPMAMUI AS ONDIFMA
SBS jyaq
Fig, 3. Surtace and bottom salinities (TDS °/ao) and the
occurrence of macroinverlebrates and fish at Stations
7 and 9 in the North Lagoon from Murch 1983 16 March
1985.
Tanviarsus. harhitarsis was easily the dominant
littoral and benthic animal and it occurred
tiroughout the South Lagoon on most occasions
although with high salinities in March 1984 (Fig. 4)
and March and April 1985 (Fig. 4) it did not occur
al Stations 6S er 7S. At Stations 2S to 3S the
abundance of 7) barhjtarsis was seasonal with
highest numbers if summer (D, Paton pers..comm,),
Six fish species were collected in the North
Lagoon (Table 1). The small mouthed hardyhead,
Atherinosoma microstama, and the blue spot goby,
Psendosobius olorum, wete the ones mast
commonly collected by hand net. A. microstoma
was distributed throughout the North Lagoon while
P. olorum was testricted to more northerly areas
until mid 1984 when salinities fell (Fig. 3). The other
four species were collected only occasionally.
ACTS me >
buicher! -
Alcrionerta
forse?
Peevcaanritis
orwall’
Atherimasomg
WUCTES HNO.
Tanytorsus
barhitarsrs
Ruppie
fufieraso
— |
7
2 oc ny Ses xe
Apri! BS
a ae
a 4 ——5__+
-24{ or Sa 1S
6 O5 636 & OF
~ e = = q
; = O° © é3 fe)
‘i £ é = #
1 4 : z
e 2 # % 2
4 v Sa
Fig, 4, Longitudinal salimty profiles and ihe distribution
of biota in the South Lagoon during a low salinity
peviod in spring (September-Ocrober) 1984 and.a high
salinity period in autumn (March-April) 1985,
A much more complete fish sampling program
was. conducted in the North Lagoon of the Coorong
during 1984 by David Hall of the S. Aust. Dept
of Fisheries. The common species in. his catches,
along with those in the present study comprise a
complete list of the common fish occurring in che
Coorong in 1984 (Table 2), They include freshwater
species presumably washed in with the River Murray
flow, marine species thal usc the Coorong as a
nursety-growing aréa, and species that generally
complete their life cycles in the estuarine-lagnonal
system.
The hardyhead, A. mucrostoma, was the only fish
that persisted in the South Lagoon. Hardyheads
occurred in large numbers throughout the South
Lagoon in winter, spring and early summer.
Increasing salinities in late summer and autumn
restricted their southerly distnbution so that only
a few specimens were collected at Station 7S in
March 1985 when the salinity was 149%/o0, and
they were still rare in April 1985 when the salinity
had fallen to 100°%co (Fig, 4), In late winter and
spring 1984 when salinities fell as Juw as 35°00,
congolli (Pseudaphritis urvill), black bream
(Acanthopagrus bulcheri) and yellow-eye mullet
(Aldrichetta forstert) also occurred in the South
Lagoon (Fig. 4). Yellow-eye mullet and black bream
were caught by professional fishermen in the
northern section of the South Lagoon in August,
September and October 1984 and recreational
catches of mullet were recorded as. tar south as Sale
Creek (D. Hall pers. comm.), No catches of
congolli, black bream or ycllow-cye mullet were
recorded in July or November when TDS was
around or above 70°%oa,
COBRONG BIOLOGY ANIT RIVER MURRAY FLOW {73
Discussion
The high River Murray outflows frou tuly-
November 1983 had little inumediate impact an the
salinities in the North and South Lagoons, In
December 1983 salintlies were-as high as 6D°/oo at
the south end of the North Lagoon and above
30°o0 ty the South Lacoon. The major drop in
salinities in rhe North Lagoon occurred between
December 1983 and Murch 1984, resulting in
salinities below 30°%oo throughout the lagoon,
During tus perlad outflows from the barrages wore
moderate. Thus mixing of fresti water soulhwards
in the Coorong is not simply controlled by River
Murray Flow but other factors such as sea levels,
lagoon levels, wind direction and evaporation are
probably Involved (Nove 1975}. Freshening of the
South Lagoan eccurred in the latter half of 1984
so that In September-October salinities were 50-
WM, These salinities were still well above those
recorded in November 1975 (30-40°%o0) following
@ period of yery heavy River Murray flow in 1973
and up lo 1975 (Geddes & Butler 1984),
The River Murray flow produced significant
effecls upon the Ruppia populations in 1983 and
1984, In the North Lagoon, Ruppia megacarpa, a
species which usually reproduces vegetatively and
sets little seed (Brock §982), lowered profusely in
October 1983 after the first heavy outflows from
the River Murray and the slight fall in salinity. In
the South Lagoon, Ruppia tuberosa became
abundant in 1984 and thick and extensive beds of
lowering plants were observed in October-
December 1984. This followed a drop in salinity to
§5-70%w along the South Lagoon in September
1984, This abundance of R. rvberosa contrasted
with Its scarcity during 1982, when salinities were
Benerally above XP /ie (Geddes & Butler 1984), and
supports observations made by others (Delroy e7 ai.
1965'+ Womersley 1975: Paton 1982+} chat growth
of R, ruberosa is inhibited at salinities above Lwice
Seawater,
The minor dilution of the North Lagoon during
1983 did not produce any changes in the distri-
bution of macroinvertebrates, However, with the
marked salinity fall in March 1984, most species
extended their distributions southwards although
there was a Jaz period with must species nat
Occurring at the mosi suutherly station until June,
' Delory, L. B., Macrow, PM. & Sorrell, J. B. (1965) The
foud of warerfow! (Anadidae) in sult water habirars of
Sourh Australia, Unpublished reporr of Fisheries and
Fauna Conservation Department of South Australia,
* Paton, P, (1982) Biotn of the Coorong. South
Australian Department af Environment and Planning,
Nov. 1962. A.DIEP. 5$ (inputlished),
No members of the estuarine-lagooual imacro-
invertebrale fauna were collected In the Sourh
Lagoon although salinities there were within their
tolerance Jimits in September-Detober 1984,
Presumably longer periods at moderate salinily are
mecessary lor the establishment of the estuarine-
Jagoonal fauna in the South Lagoon. The “sale Jake"
association of dipterans, ostracods, Halanisens
searieé and Alherinasomea nricrosioma persisted. in
the South Lagoon throughout 1983 and 1984,
The more estuarine conditions i 1983-84 did iho
see any significant increase in the number of spevie's
of macroinvertebrares inthe North Lagoon. Only
one lurther polychaete, Prionasplo cirrifera, anc
one further gastropod, 7atea rajfilabris, were
collected. Prinnoespio sp. is a commonly
encountered polychaete in the Leschenaalt and Peet-
Harvey estuaries in south-western Australia (E, P,
Hodgkin pers. comm.) and Prionespio virrifera is
common in many estuaries On the south eust
Austrahan coast, especially where there is
considerable freshwater inpul or eulruphicalion
(Rainer & Fitzhardinge 1981; Collett ey af 1984}.
Tatea rufilubris is a hydrobiid found in southern
Australian estuaries (Ponder pers, comm.)- Thus,
even during periods of River Murray flow and
moderale salinities, lhe number of species in the
Coorong (23 species of macroinvertebrates) was
much lower than those recorded in some other
Australian estuaries (Rainer & Pitzhardinge (98);
Geddes & Butler 1984),
River Murray flow and the changed salinity
pattern brought about changes in the fish fauna and
the distribucions of particular specics. The
freshwater species, Cyprinus carpie, Nemeatalosa
erebj and Philypnodon grandiceps, occurred in the
North Lagoon along with the matine and estuaiine
fish, The three freshwater species presumably were
washed in with the River Murray flow, but kirge
populations of N. erehi persisted alone (he entire
length of the North Lagoon for considerable
periods in 1983-84, Three &pecies previously
restricted to the North Lagoon, Pxeudaphritis
uevilli, Acanthepagrus butcheri, and Aldrichettu
forsteri, entered the South Lagoon for a brief period
in August-October 1984 when sallnities were below
70%oo. The disttibutions of these commercial
fishes seem to respond rapidly ta the establishment
of favourable salinities, although catch per unit
effort data suggest that only small numbers of fish
were present (D. Hall pers, comm,),
The fish fauna of the Coorong, like that of the
invertebrates, is very restricted campared fo thal in
other estuaries (Pollard 1984). This is mustly cue
ta the minor represenration of marine fish in che
Coorong. OF the 17 common species ( lable 2), only
five pre marine species that use the Coorony
18
M. C. GEDDES
Tasi_e 2. Common fisk in the North Lagoon of the Coorong during 1984. Species are grouped according ta their
habitatas follows: estuarine — can. complete lifecycte in the Caorang; marine — generally reproduce at sea; freshwater
— generally reproduce in fresh water,
Family Species nume
ESTUARINE
Atherinidac ' atherinasoma microstoma (Giinther)*
Gobiidae Pseudouobins olorum (Sauvage)
Arenogobius bifrenaius (Kucr)
Callogobius mucosus (Ginther)
Bowichyilae ' Pseudophritis -arvilll (Valenciennes)*
Sparidac Acanthopagrus buteheri (Munro)
Bothilae Rhombosolea tupirina (Ginther)
Hemiramphidae Hyporhamphus regularus (Gunther)
Engraulidac * Engrautis australis (Shaw)
MARINE
Setaenidae Arpyosomus hololeptdatus (Lacépéde)
Mugilidae Aldrichetta forsteri (Valenciennes)
Clupeidae * Sardinopy neopilchardus (Steindachner)
Galaxndae 4 Gulaxlus maculatus (Senyns)
FRESHWATER
Clupetdae Nematalose evebi (Gunther)
Eleotridae Philypnodon grandiceps (Krett)
Cyprinidae Cyprinus carpia (L.innenus)
Common name
Smal mouthed hardyhead
Blue-spot goby
Bridled goby
Sculptured goby
Congalli
Black bream
Greenback flounder
River garfish
Southern anchovy
Mulloway
Yellow-eye mullet
Australian pilchard
Common galaxias
Bony bream
Big-headed gudgeon
European carp
' These, species can breed in tresh water (Lloyd unpublished).
* Breeds in estuaries as far as is known.
* Breeds at sca as far as is known,
4 Breeds in estuaries but larvae then spend time al sea before returning to estuary of fresh water (Pollard 1971}.
lagoons as 4 nursery-growing area, while there are
perhaps nine species thal are resident within the
syste. A similar, but not so severe, restriction of
usage by marine specics has been noted for the Peel-
Harvey system (Potter ef a/, 1983) and the Swan-
Avon estuary (Prince & Potter 1983) in Western
Australia. These authors suggest that the presence
of a narrow channel and of extensive peripheral
bays within the estuaries, make these systerns
conducive to an estuarine mode of life. Conversely
these same factors make it difficult for marine fish
to move in and out of the estuarics. ‘he extreme
salinity Mluctuations in (he Coorong may also limit
lish diversity.
Hypersalinity can be a major luctor limiting fish
distribution within estuarine-lagoonal systems.
Only six fish species occurred in Hamelin Pool, the
hypersaline (~54%oo) region of Shark Bay
(Lenanton 1977), and while most species in South
African estuaries-lagoons could tolerate salinities
up to $5°%eo, only a few could tolerate salinities up
to: 70%0 (Hill 1981). A similar dimunition of fist
fauna between 55 and 70"%/oa appears to occur in
the Coorong, The most tolerant species in fhe Shark
Bay aod South African studies included members
of the Atherinidae, the Mugilidae and the Sparidae,
families which also include the most salt tolerant
species in the Coorong. Considenng the effects of
salinity on the distributions of fishes within the
Coorong, it appears that under the present hydro-
logical and salinity regimes the fish faunas will
sometimes be restricted in the southern end of the
North Lagoon and on most occasions only the
highly tolerant Athkerinosume microstoma will
occur in the South Lagoon,
The effects of the River Murray flow of 1983-84
on salinity patterns and rhe distribution of biota
in the Coorong were short-lived, Following the
barrage closure in December 1984, salinities quickly
rose to 36-70%e0 in the North Lagoon and
80-140%50 in the South Lagoon by March 1985,
This is an almost identical salinity pattern to that
of March 1983 afer a period of extended barrage
closure. It seems that consistently high River Murray
flows are needed to maintain an estuarine-marine
situation in the North Lageon and moderately
hypermarine conditions in the South Lagoon of the
Coorong.
COORONG BIOLOGY AND RIVER MURRAY FLOW 141
Acknowledgments
Thanks to Helen Vanderwoude, David Paton,
Greg Powell and Julie Francis for help in the field
and the laboratory and the following people for
identifying specimens; Pat Hutchings (polychaetes),
Jerry Barnard (amphipods), Winston Ponder
(gastropods) and Lance Lloyd and David Hall
(fish). David Paton and David Hall also allowed
access to unpublished data. The Engineering and
Water Supply Department provided unpublished
data on outflow from the River Murray Barrages
and the South Australian Fisheries Department
provided data on fish species and disiribulion,
Thanks to Sandra Lawson for typing the manu-
scnpt, to Ruth Evans for the artwork and to the
Zoology Department, University of Adelaide for
support.
References
Brock, M. A, (!982) Biology of the salinity tolerant
genus Ruppia L, in saline Jakes in South Australia. II.
Population ecology and reproductive biglogy. Aquat.
Bot. 13, 249-268.
CouiceTT, L. C., AuTcHmas, P. A, Gras, P. J, &
Coins, A. J. (1984) A comparative study of the
macro-benthic fauna of Posidonia australis seagrass
meadows in New South Wales. Aquat. Ror, 18, 111-134.
Geopes, M. C. (1984) Limnology of Lake Alexandrina,
River Murray, South Ausiralia, and the effects of
nutrients and light on the phytoplankton. use J. Mar.
Freshwat, Res. 35, 399-415,
& Butter, A. J. (1984) Physicochemical and
biological studies on the Coorong lagoons, South
Australia, and the effect of salinity on (he distribution
of the macrobenthos. Trans. R. Soe. S$, Aust. 108, 51-62.
Hii, B. J. (1981) Adaptations to‘temperature and salinity
stress in South African estuaries, Ch. 11, pp, 187-197,
In J. A, Day (Ed.) “Estuarine Ecotogy”. (Balkema,
Rotterdam).
HurtcHinas, P. A. & Giasuy, C. J, (1985) Addirional
nereidids {Polychacta) from eastern Avistralia, together
with a redescription of Nammanerets quadraticeps (Gay)
and the synonymising of Ceratonereis pseudoery-
thraeensis Hutchings & Turvey with C. aeguisetis
(Augener), Rec, Aust, Mus. 37(2), 101-110,
Kanaas, M. L. & Gepprs, M. C. (1984) The effects of
salinity on the distribution of amphipods i the
Coorong, South Australia, in relation to their salinity
tolerance, Trans. R. Soc. S. Aust. 108, 139-145.
LenanTon, R. ©. J. (1977) Fishes from the hypersaline
waters of the stromatolile zone of Shark Bay, Western
Australia. Copeia (1977), 387-390.
Noyt, B. J. (1975) The Coorong. Publ. No. 39, 131 pp.
(Department of Adult Education, University of
Adelaide).
POLLARD, D. A. (1971) The biology of a landlocked form
of the normally catadromous salmoniform fish Galaxtas
maculatus, 1, Life cycle and origin, Aust, J Mar
Freshwat. Res. 22, 91-123,
(1984) A review-of ecological siudies on sea-grass
fish communities, with particular reference to recent
studies in Australia. Aquat Bol. 18, 3-42.
Porver, I. C., Loweracan, N, R, LRNANTON, R. Cb,
Curysta, P. J. & Grant, C. J. (1983) Abundance,
distribution and age structure of fish populations in a
Western Australian estuary. J. Zool, Lond. 200, 21-50.
Prince, J. D. & Potter, 1. C. (1983) Life-cycle duration,
growth and spawning times of five species of Atherini-
dae (Teleostel) found in a Western Australian estuary.
Aust, J. Mar, Freshwar. Res. 34, 247-301,
Rainer, S&S. & FitzHarpince, R, C, (1981) Benthic
communities in an estuary with periodic deoxygenation,
thid. 32, 277-343.
Womers.ey, H, B. S. (1975) The plants. Ch. 7. Jn
J. Noye (Ed.) “The Coorong”, Publ, No. 39 (Department
of Adult Education, University of Adelaide),
SALINA BED INSTABILITY AND GEODETIC STUDIES AT LAKE EYRE,
SOUTH AUSTRALIA
by J, A. DULHUNTY*
Surmary
DuLaustyY, J. A, (1987) Salina hed instability and geodetic studies at Lake Eyre, South Australia. Trans,
R. Soc. S. Aust 1V1(4), 183-188, 30 November, 1987.
Horizontal compressional forces in saltcrusts of Lake Eyre cause fracturing with overthrusing in chin
crusts oF marginal areas, and warping or buckling in central areas where crusts are thicker and stronger,
Uneonsolidated Holucene sediments adjust by deformation to warping of overlying saltcrusts and the true
equilibrium level of the lakebed surface is elevated or depressed, When the lake is filled salicrusts are dissalverd
and deforming pressures are released, allowing the surtace of the sediments to return towards equilibrium
level under gravity, Claims that any one place, in any one bay, is the lowest place in Australia, are doubtful,
A more appropriate concept may be that the lowest landsurfaces on the Australian continent are in the
southern bays of Lake Eyre North.
Kry Worbs; Lake Eyre, salterusts, lateral forces, cverthrusting, warping.
Introduction
Lake Eyre ts a Jarge arid ephemeral terniinal lake
Uonns 1955; Bonython 1955, 1956; Dulhunty R.
1975, 1984, 1986; Dulhunty JA. 1977, 1978, 1982:
Allan ev af. 1986; Callen & Wells 1986), It is the
sump of an internal drainage basin, consisting
mostly of semiarid and arid country covering
almost one sixth of the Australian continent, River
courses draining the Lake Eyre Basin seldom carry
flowing waler as far as the lake, as it is mostly lost
to evaporation in desert country of low relief
through which it must pass. Small amounts of water
which do reach the lake, at intervals of 2 to 3 years,
cover only parts of the bed and usually dey up in
less than one year, Infrequent major fillings, once
in 25 to 50 years, cover the whole of the bed to
depths of § to 10 m,-and may require 4 to 6 years
io. dry up (Bye ef al, 1978; Dulhuntry R, 1984; Allan
1985},
Lake Eyre lies in the most arid region of
Australia, with an average annual rainfall of less
than 127 mm, maximum summer temperarures up
‘to #lPC (Price 1955), and an annual evaporation
rate of about 2.5 m (Bonython 1955; Penman 1955).
The purpose of this paper is to record evidence
of lakebed instability or warping of saltcrusts and
deformation of Holocene sedimenis in the salina
arca of Lake Eyre North, and its significance in
peodetic studies and the concept of the lowest place
an the Australian continent.
Saltcrusts
Saltcrusts up to about 30. em in thickness occur
in the three southern bays ol’ Lake Eyre North,
* Department of Geology and Geophysics, Universitv ol
Sydney, N.SW. 2006.
where final evaporation of brines takes place. The
salt has been transported into Lake Eyre. dissolved
in riverwater aud groundwater, after the lake
became terminal following the onset of late
Pleistocene aridity which has persisted through
Holocene to the present day (Bowler 1978;
Dulhunty J. A, 1982). When sufficient water enters
the salina area during major fillings, pre-existing
saltcrusts are dissolved; new sediments are
deposited, and finally, with evaporation of water,
salt is reformed into new crusts (Dulbunty J, A.
1982),
Three generations of gsaltcrusts have been
tecorded at Lake Eyre, The first occurred prior to
the [949-50 filling (Madigan 1930); the second
occurred between the 1949-50 and 1974 fillings
(Dulhunty J. A. 1974, Dulhunty R. 1975; Bonython
1956; Johns 1963), and the third formed after the
water of the 1974 filling dried up in about 1979
(Dulhunty R, 1984). In addition to the three
recorded crusts, a long series OF prehistoric saltcrusts
of gradually increasing volume, must have existed
between major fillings in fate Pleistocene and
Holocene times.
In 1972 @ survey of saltcrust thicknesses was
carried out by Dulhunty over Madigan Gulf and
Jackboot ard Belt Bays, The publication of results
(Dulhunty J. A. 1974) included records of crust
thicknesses measured previously in Madigan Gulf
by Madigan (1930); Bonython (1956), and the South
Australian Geological Survey (Johns 1963).
Differences in salt thicknesses measured at different
times, in the same places in Madigan Gulf, were
also recorded and discussed.
Levelling surveys of thé lakebed surface at the
base of the saltcrust in Madigan Gulf were made
in 1954 by Bonython (1956); in 1969 by the South
Rd JA. DULHUNTY
Australian Fands Department (Simmons 1969)!
and in 1972 by Dulhunty J. A. (1974). The upper
surface af rhe sediments at the base of the saltcrust
is regarded as the present stratigraphical suriace of
(he lakebed (Bonython 1956; Dulhunty J. A. 1974).
The overlying salterusts arc removed perindically
by solution during major fillings, and when
teformed after evaporation of water, they occupy
a younger statigraphical horizon. Therefore salt-
crusts cannot be regarded as specific strata, ar dated
as such, Differences in level values of the lakebed
surface at different times in the same places over
a total period of 18 years emerged from the three
above surveys during the life of the one saltcrust,
When results of both lakebed levelling and salt-
crust thickness surveys were considered in relation
to ime and place, it was evident that there had been
“normal” changes 1 thickness which occur as crusts
nature, Yesulting in migration of salt from higher
marginal areas 10 lower central regions of the
shallow saucer-shaped bottoms of the bays, as
described by Bonython (1956), In addition, however,
there appeared to be evidence of anvther lactor
Which had influenced changes in crust thickness and
fakebed surface levels during the life of the crast
hetween [he 1949-50 and 1974 fillings. This
operated independently al “normal” thickness
changes in marginal and central positions, It
elevated Or depressed relatively small areas of
salicrust and Underlying lakebed surface, which lead
to thinning or thickening of the crust respectively,
by migration of salt as illustrated in Pig. 2-
Lakebed instability
Subsequent research between 1978 and 1983,
during the formation of a new saltcrust following
the drying up of the 1974 filling, has provided a
clearer understanding of the factors mvalved in
what may now he described as lakebed instability.
A series of lakebed profiles hased on results of
crust thicknesses and lakebed surface levels, has
been consinicied in Fig. 1 to illustrate lakebed
instabilicy within a limited area in (he central region
of Madigan Gulf. Section LA af Tig, 1 is a map
showing puints A lo J at which it was possible lo
compare crust thicknesses and lakebed surface levels
at different times during the life of the crus! fron
about 195] to 1974, helween the 1949-50 and 1974
fillings of the lake. Section 1B of Fig. 1 shows twa
lakebed profiles along the line ABCDEFG with sall-
crust thicknesst's and Jakebed surface levels based
on surveys by Bonython (1956) in 1954 when the
1Simmons, P RB (1969) Measurement of tesr range for
Iuser ARP at Lake Eyre, South Australia Pands
Dopartment, Adelalde (unpublished),
crust Was about | year old, and Dulhunty (1974)
in 1972 when it was About 19 years old. The dif,
ferences in formes of the profiles show the amounts
of change in lakebed surface levels between the two
surveys. Also, crust thicknesses indicated by S values
along the profiles show the amounts Of change in
crust thicknesses which had occurred between the
two surveys.
Section tC of Fig. | shows lakebed profiles along
the line 1/13, in 1954 and 1972. Section 1D of Fig.
| shows three profiles with level values along IHG.
Two of these have crust thicknesses indicated by 5
values based on surveys in 1954 and 1972, The chled
profile withoue crust thicknesses in Section 1D, is
hased on the level survey by che Sourh Australian
Lands Department in 1969, during which na crust
thicknesses were reported. However, the relatively
Yow level value of this profile in 1969, in relation
wo the higher values of earlier and Jater surveys.
indicates the complex nature of changes in level
values that had occurred teiween 1954 and (972,
Where salicrust and lakebed surlace rise Over an
upward buckle, the crust becomes thinner by trans-
fer of salt to a Jowee area such as a nearby down-
ward buckle as illustrated in Pig- 2, B and C.
Conversely, when the crust and lakebed surface sink
over a downward buckle, the crust thickens by
iransfer of salt from a nearby higher area, As well
as thickness changes due to buckling, and super
iinposed upon rhein, “normal” wansfers of sale frou
higher marginal areas to lawér central areas, take
place during the life of the crust, Thus thicknesses
in a buckled arca are the net resutt of boeh factors.
In the buckled areas ilustnited in Figure | there are
8 points at which lakebed surface levels and crust
thicknesses. were hoth determined by Boaython in
1954 and Duthunty in 1972, Of these & points, aver
the perind of FS years, 34AB & () went up and
crust thinned and 4 (GHI& J) went. down and enust
thickened. At only | point (D) had the net crust
Ihickness not conformed to thinning with rise and
thickening with fall, At this poimt addition of salt
due to some other factor such 8s mignition from
marginal to cenwal areas, or a nearby rise on an
upward buckle, mush have heen greater than loss
of sale due to rise in Jakebed surface level.
It is now concluded that lateral compressional
forces, generated in sattemuists as they marure, are
responsible for lakebed instability, Evidenve of the
operation of horizonta) compressional forces in thin
and relatively weak saltcrusts up to about 10 em
thick, in marginal areas, is seen in extensive
fracturing of the crust with overthrust movement,
illustrated hy photographs in Fig. $. The berzontal
displavement associated with overihrusting on
individual fractures is as much 30 cm in same
places, but wsually less than 1S em, There is a great
SALINA STUDIES AT LAKE EYRE 185
MAP
CENTRAL AREA
MADIGAN GULF
Ben IN LAKE EYRE NORTH
CAMPBELL SHOWING
LEVEL SURVEY POINTS
NSULA
HUNT PENI
1
ey
>
=)
___
HORIZ. SCALE
re)
Salt 31cm
DULHUNTY 1972
elow SL.
Aust. Hgt, Dtm.
Oo
e 1
Salt 27.9.,, $27-9 §27-9 $279
0 2
HORIZ. SCALE
Salt 28cm
DULHUNTY 1972
BONYTHON 1954
J
iy 2 4km
§22 HON 1954
HORIZ. SCALE BONYTHON 195
23 $23
$42
s
Oo DULHUNTY 1972
Salt33.cm
$35
tt LANDS DEPT. 1969
H G
FIG. 1D caxeseo proFies
Fig. | Lakebed surface profiles and saltcrust thickness along lines in central Madigan Gulf of Lake Eyre North.
186 I, A, BULHUNTY
EEO IMENTS
“Tavexteo
SALT AMD
SURFACE
OF BED
SEDIMENTS
TELUTM WZ
eg ee
5
SAU
TRANSFER
6Y
[one ION
pal som NING
DEPOSIT
AND
DEPOSITION
Nig. 2. A — diagranumatic WMustration of surface of unconsolidated Holocene sediments beneath salicrust. B— buckling
of salicrust and underlying sediments by lateral compressional! farces. C — sali transfer From higher to lower place
by solution and redeposition of salt giving thinning and thickening of crust
deal of fracturing in all marginal areas surrounding
the principal sheets of saltcrust, as well as in small
areas formed in washouts which occur in tiver
estuaries and form walerholes after Mowing ceases
(Wopfner H. (982, pers. comm).
In central areas of the main saltsheets, where
<rusts are 30 to 50 em thick, and stronger, they do
not fracture bui warp either up or down, above or
below the true or equilibrium lakebed surface level
under gravity, by as much as 0.5 m, Warping may
occur over limited areas up to several kilometres in
width, The watery pliable nature of the underlying,
unconsolidated Holocene sediments allows adjust-
ment by deformation to the warping of the more
rigid saltcrusts, und so the level of the lakebed
surface is elevated ot depressed,
Fracturing, with up to 10cm of horizontal over.
thrust movement, occurs in small isolated areas of
crust, from 100 to 300 m across, such as those
formed in washouts along river estuaries. This
suggests that appreciable amounts of compressional
force, or 6xpansion of the crust, are generated
throughout both marginal and central regions of
the large saltshéets in the southern bays of Lake
Eyre North, Also, it suggests that fracturing and
buckling in marginal and central regions is due 1a
compressional dorces gencrated within those
regions, and not necessarily transmitted long
distances from one region to another across the
saltsheet.
In addition to well-defined fracturing of thin
crusts with overthrust movement, and warping of
thick crusts, a polygonal jointing pattern is
developed in saltcrusts. Brine rises from ground-
water along the cracks to the surface, where it dries
and deposits new crystalline salt. 1f the crust is very
thin (less than about 7 cm) new salt may be
deposited along cracks beneath the polygons, pro-
ducing upturned edges around the slabs which vary
from about { to 4m in diameter. Where the salterusc
is thicker (10 cm or more) the new salt forms low
ridges up to 5 cm high over the cracks, but it is
eventually dissolved in rainwater which dries and
the salt is added to the pre-existing crust (Madigan
1930; Bonython 1956; Johns 1963).
It has been suggested by Prof, C C, von der
Borch (1986, pers. comm.) that the fracruring,
warping and polygonal jointing described in this
paper may be analogous to processes which produce
“tepee” slructures (Assereto & Kendall 1977; Reeves
1970) which commonly occur in expanding
supratidal carbonate crusts analogous to expanding
halite crusts. This is very probably so, but the actual
physical or chemical mechanism in the origin of
tepee structures in relatively short-lived salt crusts,
is a complex matter well worth further investigation,
FLORICEPS MINACANTRUS SP. NOV. (CESTODA: TRYPANORHYNCHA)
FROM AUSTRALIAN FISHES
BY R. A. CAMPBELL* & I. BEVERIDGET
Summary
Floriceps minacanthus sp. nov. is described from adult worms in sharks and from plerocerci
obtained from the viscera of teleosts taken in Australian coastal waters off Queensland, New South
Wales and South Australia. Selachian hosts are Carcharhinus brachyurus (Guenther, 1870)
(type host), and C. amboinensis (Mueller & Henle, 1841). Plerocerci were obtained from the
teleosts Platycephalus laevigatus Cuvier, 1829, Plectropomus ? leopardus (Lacépéde, 1802) and
Sphyraena novaehollandiae Guenther, 1860. Descriptions of the adult and metacestode stages are
provided. Characteristics of both the adult worms and plerocerci are consistent, with the exception
that the scolex of adults is more elongated than plerocerci thereby increasing the ratio of bulb length
to pars vaginalis. Characters differentiating F. minacanthus from other species of Floriceps are the
shape of principal hooks 1(1’), shorter bulbs (<O0.9mm), ratio of pars bulbosa to pars vaginalis
(almost 1:6) and genital pore in posterior '/, of segment.
KEY WORDS: Floriceps, Trypanorhyncha, new species, Cestoda.
FLORICEPS MINACANTAUS SP. NOV. (CESTODA: TRYPANORHYNCHA) FROM
AUSTRALIAN FISHES
by R. A. CAMPBELL* & J, BEVERIDGET
Summary
Campnrbiy, R- A & BevesipGe, |, (1987) Floricens mingcanrhus sp. nov. (Cestoda: Trypanorhyncha}
from Australian fishes, Trans. R. Sev, S. Aust. 114(4), 189-194, 30 November, t9&7.
Flariceps minacantius sp. nov, ss described from adult worms in sharks and from plerocerci obruined
frorn the viscera of teleosts taker in Australian coastal waters off Queensland, New South Wales and South
Australia, Scluchian hosts are Carcharhinus brachyerus (Guenther, 1870) (type host), and C. anzhoinenyis
(Mueller & Henle, 1841). Plerocerci were obtained fram the teleusts Platycephalus laevigatus Cuvier, 1829,
Plectropomus ? leopardus (Lacépéde, 1802) and Sphyraena noveehollandiae Guenther, 1860, Descriptions
of the adult arid metacestode stages are provided. Characteristics of both the adult worms and pleracerci
are consistent, with the exception that the scolex of adults is more elongated than pleracerci thereby increasing
the ratio of bulb length to pars vaginalis, Characters differentiating A ryinucanthus trom other species
of Floriceps are the shape of principal hooks 1('), shorter bulbs (<0.9 mm), ratio of pars bulbosa to
pars vaginalis (almost 1:6) and genital pore in posterior '4 of segment.
Key Worns: Flarfcepes, ‘Irypanorbyncha, new species, Cestoda,
Introduction
Specics of the trypanorhynch cestode genus
Floriveps Cuvier, 1817 are common parasites of the
spiral valves of sharks in many regions of the world
(Dollfus 1942) with the metacestode stages (plero-
cerci) occurring encapsulated in the viscera of
leleosts. The genus has not previously been reported
from Tish in Australian coastal waters (Beumer et
al 1982). Recent cestode collections from sharks
and teleasts contain both adults and plerocerci of
a ew species Of Fluriveps and \he description of
the new species forms the basis of this paper.
Materials and Metis
Adult cestodes were removed from the spiral
valves of sharks and were either killed by relaxing
in Lap. water and fixing in 10% formal saline, or were
lixed {ive with hor 10% formalin, Plerocerci were
fixed in 10% formalin or the scoleces were dissecred
free, the tentacles everted by placing in fresh waler,
and they were then fixed in formalin or Berland’s
fluid. Whole mounts were stained with eclestine
blue, dehydrated in ethanol, cleared in clove oil and
mounted in balsam. Teotacles were dissected free
from scoleces, and were mounted in glycerine jelly.
Measurements are presented in the text in micro-
* Department of Biology, Southeastern Massachusetts
University, North Dartmouth, Massachusetts, USA
U2747,
| South Australian Department of Agriculrure, Ceniral
Veterinary Laboratories, c~ Mnaritute of Medical and
set Seiéiee, Frome Read, Adelaide, S Aust,
metres, unless otherwise stated, as the range
followed by the mean in parentheses. The number of
measurements mad¢ is shown in the form n=.
Terminology for typanorhynch morphology
follows that of Dollfus (1942). Specimens have been
deposited in the South Australian Museum (SAM),
Adelaide, the British Museum (Natural History)
{BMNH), London, the United States National
Museum Helminth Collection (USNMHC},
Washington, and the Australian Helminth Collec-
tion (AHC), in SAM.
Floriveps minacanthus sp, nov-
FIGS I-17
Description: Measurements of eight adult
speciinens from Carcharhinus brachyurus (types).
Scolex and strobila slightly craspedote, strobila up
to 18.5 cm long, segments hyperapolytic., Scoles
length 3550-5950 (4770) (n=8), width at bulbs
800-1470 (1060) (n-8), Two bothridia, heart-
shaped, cach with small indentation on posterior
margin, length 930-1140 (1030) (n=8) by 1300;
anterior margins curled toward midline (Fig. 15},
Pat's vaginalis 2860-5500 (4050) (n=8), tentacle
sheaths coiled; gland cells scattered in parenchyma
of pendunculus scolecis, Bulbs 680-800 (720) (1 =8)
long, 140-250 (200) (n=8) in diameter; retractor
muscle attached near posterior extremity, Pars post-
bulbosa lacking. Ratio of pars bulbosa to pars
vaginalis 1:4,2 to 1:7.3 (1:5.6) (n=8). Metabasal
armature pocciloacanthous, with single chainette
of V-shaped hooks; no basal armature. Metabasal
armature consists of altemating half spiral rows of
7 hooks each; single satellite hook Jocated posteniur
to seventh hook of each principal row (Fig, 3),
k. A, CAMPBELL & L BEVERIDGE
19)
a
lh
yi
j«
phy
KK
—<2L ~BY~B~9 OD _ ~s ap
I LBD
~ 44s S92 ALIA
we
Figs |-I4, Mloriceps minacantaus sp. nov, |, 2, metabasal armature, bothridial surface; 3, basal and metabasal armature
of tentacle, external surfave; 4, metabasal armature, internal surface; $-11, profiles of hooks 1, 2, 3, 4, 5, 6 and
7 respectively; 12, profile of satellite hook or hook 8; 13, chainette element from mid-tentacular region; 14, chainette
element from metabasal region. Scale lines: figs !-4, 0.1 mm; figs 5-14, 0.01 mm. Legend: bothridial hooks
Te: Pare 7; antibothridial hooks 1,3’, ..,. 7'; satellite haoks §; chainette C.
NEW SPECIES OF FLORICEPS 191
Hooks hollow; hooks of principal rows and
chainette smallest in basal region, reaching
maximum size 8 to 10 rows from base, decreasing
in size beyond midlength of tentacle, Hooks Lit’)
and 2(2') on inner face large, thorn-shaped; Lit'}
strongly cucved toward tentacle, leneth 21-30 (28}
(n<10), base Jength 158-20 (19) (n=10), maximum
height 10-25 (18) (n=10); hooks 2¢2') erect, nocurved
at tip, length 24-31 (29) (n=10), base 18-20 (19)
(n=10), height 15-20 (17), Hooks 3{3’) to 7({7')
sinuous, becoming spiniform in shape as row
reaches external face; 343°), Jemgth 25-32 (30)
(n=10), base length LU-14 (12) (n=10); 4(4') length
22-32 (29) (n-10), base length 8-11 (10) (n=10);
5(5") length 16-24 (22) (n =10), base length 5=8 (7)
(n=10); 6’) spiniform, length 16-22 (20) (n=10),
hase length 3-6 (5) (n=10 7(7') spiniform, length
23-30 (28) (n=10), hase length 4-6 (5.2) (n=I0),
Satellite hook (hooks 8(8')) spiniform, length
18-26 (24) (n=10), base length 5-K (6) (a=IU).
Chainetfe single, consisting of large fooks with
basal winglike processes; maximum dimensions at
base of tentacle, length 6-10 (8) (n=10) by 12-14 (13)
(n=10); in metabasal region, width across wings
24-30 (29) (9 =10) by 17.5-20 (18) (n=10). Tentacle
diameter, 70-80 (75) (qn =4) at base, 20-25 (21) (n=4)
at midlength.
Number of segments 69-83 (n=3). Segments
appear 6,0-9,5 mm posterior to scolex, First
segments wider than long, rapidly becoming longer
than wide wilh maturity, Mature segments 4200-
6850 (5070) (n=10) by 1400-1700 (1450) (n=10). Testes
medullary, spherical, 50-65 (60) (n=L0) in diameter,
filling all available space between lorgirudinal
osmorezulalory canals and extremities of segment,
icluding postovarian region, Tesles number about
1200 per segment. Genital pores marginal, irre.
gularly alternating, posterior, located 69-79% (73)
(n=10) of segment length from anterior margin.
Cirrus sac 430-3570 (S00) (n =10) by 200-300 (240)
(n=10), containing internal seminal vesicle. External
seminal vestclé absent. Vas deferens coils medially
then posteriorly fo level of ovary. Vagina posterior
to cirrus sac, Ovary bilobed, maximum dimensions
180-260 (190) (n=10) by 200-300 (270) {a=10).
Mehlis’ gland ventral to ovarian isthmus, c 50 in
diameter. Vitellaria follicular, forming a layer
enclosing osmorceularory canals and reproductive
negatis. Literus simple, median, extending ahout
three-fourths of segment length.
Measurements of 7 specimens from Carclunhinus
arnboinensis: Scolex 3250-4500 (3870) (n=7),
maxinium width 600-1200 (820) (n—6); pars both-
rstialis 710-960 (870) (fn =6); width of bothridia 750;
pars vaginalis 2300-3340 (2930) (n=7); bulbs
650-410 (810) (n=) Long by 180-290 (220) (n= 6)
if chyameler; ralio of bulbs to pars vaginalis
12.9-1;4.5 (1:36) (n=7).
Plerocerci: 28 plerocerci obtained from. viscera
and body cavity of three specics of teluost Fishes.
Armature and scolex features agree with adult
worms from sharks. Three-specimens fram Plectro-
pontus measured: Scolex 3000-3600 (3230) by
$20-1380 (1100); bothridial length 930-1040 (970):
pars vaginalis 2330-2700 (2520); bulbs 750-840
(81M) dy 200-250 (230), ratio of bulbs to pars
vaginalis 1;2,8 to :3,6 (1:3.2); tentacle diameter 70
(base), 40-60 (midlength). Eight plerocerci obtamed
from Platycephalus laevigatus; measurement of 3
specimens with extended tentacles: Scolex
4320-5000 (4680) by 850-1200 (1030); bothridial
length 1000-1060 (1040); pars vaginalis 3470-4050
(3740) bulbs 750-830 (800) by 140 to 220 (190);
ratio bulbs to pars vaginalis 14.2-1:5.4 (4.7);
lentacle diameter 70 (base), 40 (midlenstn). Single
plerocercus from Sphyreena novachallandiae
Guenther, 1860: Scolex 4760; bothridial Iengeth 1010;
pars vaginalis 3050; bulbs 775-800 by 170-215; ratio
bulbs to pars vaginalis 1:3.8; tentacle diameter 78
{base}, 60 (midlength).
Hasts and focalities; adulis; Cercherhinus
brachyuras (Guenther, 1870) (type bost), Tachra,
NSW. (lype locality) (5 specimens); Port Lincoln,
S. Aust, (4 specimens). C ambojnensis (Mueller &
Henle, 1441}, St Lawrence, Qld. (7 speciniens) (AHC
$2652),
Ste in host; Spiral valve,
Types: Holotype SAM V4UI5, 2 paratypes SAM
V 4036-4037; 2 paratypes USNMHE€ 79545, 795467
4 paratypes AHC 82650, 8265).
Pleracerei. From Plectropomus? leopards
(Lacépéde, 1802), Heron Island, Qld (3 specimens)
(AHC $2653); Platycephtalus laevigetus (Cuyiwr
1829), Northhayen, & Aust, (8 specimens) (SAM
¥4038) (BMNH 1986.10.14.2-3); Platyoephatus sp.
from fish shop, Adelaide, S. Aust. (16 specimens)
(ARC $251}, 2512, 2513); Sphynaena novachollan-
diae Guenther, 1860, Northhaven, S. Aust,
(L specimen) (BMNH 1986,10,34.1).
Site i host; serosa of viscera and in body vavety.
Etymology: The specific nanse is derived from
Latin, sinax, meaning “strongly recurved" and
relérs ta [he shape of hooks i(f').
Discussion
Schmidt (1986) listed four species helanging to
the genus Floriceps, namely the lype species, &
saccaties Cuvier, IIT, Fl cataieroy Cruz-Reyes, L977,
F Nichive Pinter, 1929 and & txvneri Guiari, 1938,
192 R. A. CAMPBELL & |. BEVERIDGE
Figs 15-17. Floriceps minacanthus sp. nov, 15, scolex; 16, mature proglottis; (7, cirrus sac and distal vapina. Scale
lines: figs 15, 16, 1,0 mm; fig, 17, 0.1 mm.
NEW SPECIES OF "LORICEPS 193
F, savoatus ditters from & ininacnarhaus sp. nov.
in faving hooks ((1') and 2(2') of Lhe same shape
rather than with hooks 1(1') almost recumbent,
larger bulbs (3 nim or more in length), pars bulbosa
to pars Vaeitalis ratin of fess than 1:3, and bothridia
that are more triangular in shape In comtrast haoks
11 of A mtiinecsnithus ts extremely arched and
almuyst recumbent whereas hooks 2(2') siand ereci
as in F saccatis, the bulbs are short (S40-or less),
the ratio of pats biulbosa ro pars vaginalis averages
1:5.6, and the bothridia are more rounded in shape.
Gland cells within the pedunculus scolecis extend
froin the bothridia to the antenor end of the bulbs
jn # minacanthus, while in KR saceafus they are
sicher restricted to the antenor part of the pendun-
culus scolecis (Dollfus 1942, figs 198, 200), or are
sancentrated in this region (Linton 1924, fig. 1, as
Ripncobothriun ingens).
A series uf characters present in the mature
segment may also serve to separate the two species.
Yamaguti (1934) estimated that in # saecatus (as
Dasyrivnchus ingens (Linton}}, 200 testes were
present in a single sagittal section, compared with
about 1200 per segment found in F mrinucunfhis.
Even allowing for several layers of testes in A
saccoius, testis nuniber may differ between the rwo
species, In minecanthus, an internal Dut not an
external seminal vesicle is present (Fig. 17), whereas
both Linton (1921) and Yamaguti (1934) describe
both vesicles as being present. linton (1921) has the
two vesicles incorrectly labelled in his figures (A and
12), while Yarnaguri’s figures of the external seminal
vesicle are not convincing, This feature of the
anatomy of F saceatus should be re-investigvated as
it muy be a further characier separating the wo
species.
All reports of F sacewtus in which measurements
are given, summarised hy Dollfus (1942), agree in
having bulbs approximately 3.0 mm long, Sub-
hapradha (1955) however, reported “A saccuits"
from Carcharhinus sp. from Madras, in which the
bulbs were 0.75 mm Jong and hooks 1(1') are clearly
drawn as being recurved (Figs I3b, 14). This
specimen is considered to he /) minacanthus and
considerably extends the geographic range of the
mew species,
Crug-Reyes (1977) provided a single view of ihe
armature of F cabalferer and his figure of the
chainetre (4a) shows hooks quite uolike those of
Fo minucanikus. The correct orientation and detail
of the armature of F c@ballerai is quite unclear
from his description and al needs WW be redescnbed
before any critical assessment of its status can be
made. FA caballerol can be distinguished from
F minacanthus becuuse the bulbs of & caballeroi
are long (3 mm), Cruz-Reyes (1977) described the
mature segment of Fcadisferal as having (he genital
hores ih the anterior fourth of the segment, the
vagina passing anterior to the cirrus sac, the uterus
terminating ar the level of the cirrus sac and the
lestes being numerous chaush few are shown in his
drawing (Fig, 5), leis highly unlikely that the mature
segment described by Cruz-Reyes (1977), which is
in fact a detached segment, is that of a crypano-
rhynch.
Dullfus (1942; p. 395) noted that F fichiae of
Pintner (|929) is very similar to Moficela horrida
(Goodsir, 1841). This means thay Pintner’s species
possessed four bathvidia unlike species of Flariceps
which have only two (see Pintner 1929). Guiart’s
(1938) description of # axnert ts very poor, such
that it cannot be identified to any specific trypano-
rhyoch family. The two microvraphs of the plero-
cetus show no detail of the bothridia or armature.
We consider F oxneri as incertae sedis.
Other trypanorhynchs chat have been wrongly
assigned to Floriceps are FD suncinatus (Linton,
1924), F crassicolle (Diesing, 1850), F elongaries
(Rudolphi, 1819), & macmcercus (Rudalphi, 1819),
and F granvius (Rudolphi, 1819). Yamaguti’s (1932)
descriptiun and figures of F wrcinatus appear to
beaspoues of Gyrrmarfprchus though he does not
list the species in his subsequent synopsis of the
trypanorhynchs (1959). According to Dollfus (1942)
all of che remainder belong in the Gymnorhynehi-
dae except F granuius which is a synonym of
Lacisterhynehus tenuis (Beneden, 1858).
The only major dilference belween adults and
plerocerci of FF minacanthus is the increase in soolex
length of the adult worms as compared to ihat of
the plerocerci. The increased length affects the
length of the pars vaginalis and accounts fol the
difference in the ratios of bulb length to pars
vaginalis. However, the bulbs are the same size in
both the pleracerci and adults and arc the smallest
bulbs of any species in the genus Floriceps.
The hook numbering system used bY Dollfus
(1942) for F swcevties has been utilised in the above
description. However, the so-called “sateflite hooks”,
which he ammediately postenur to hooks 7(7') in
Fforiceps occur in an exactly analogous position in
Cotlitetrarhvachus Pintner, 193f and Lacisto-
réprchus Pintner, 1913, in which genera they wre
simply desvribed as hooks &(8'), There is camplere
homology between the hook arrangemenrs uf the
metabasal armature of these three genera, and this
homology could be stressed by the utilisation of a
uniform system of hook numbering.
Acknowledgments
Thanks are due to BG. Robertson, D. M, Spratt,
R, G, Lester, R, A. Bray, K. 1, Gowletl- Holmes and
194 R. A. CAMPBELL & 1, BEVERIDGE
D. C. Lee for the collection and provision of
specimens described in this paper. This work was
supported financially by the Australian Biological
Resources Survey.
References
Beumer, J. P., ASHBURNER, L. D,, BurBuRY, M. E.,
JeTTE, E. & LATHAM, D. J. (1982) A checklist of the
parasites of fishes from Australia and its adjacent
Antarctic territories. Technical Communication no. 48
of the Commonwealth Institute of Parasitology,
Commonwealth Agricultural Bureaux, Farnham Royal,
pp. 99.
Cruz-Reyes, A. (1977) Céstodes de peces de México, I.
Descripcién de una nueva especie del género Floriceps
Cuvier, 1817 (Trypanorhyncha: Dasyrhynchidae Dollfus,
1935). Publicaciones Especiaties Instituto de Biologia,
Universidad Nacional Autonoma de Mexico 4, 343-355,
DoLirus, R. Ph, (1942) Etudes critiques sur les tétra-
rhynques du Muséum de Paris. Arch. Mus. natl. Hist.
nat, Paris 19, 1-466,
Guiart, J. (1938) Etude parasitologique et épidémio-
logique de quelques poissons de mer. Bull. Inst.
Océanogr. Monaco No. 755, 1-15,
Linton, E, (1921) RAynchobothrium ingens spec. nov. a
parasite of the dusky shark (Carcharhinus obscurus).
J. Parasitol. 8, 22-32,
PINTNER, T, (1929) Studien ueber Tetrarhynchen nebst
Beobachtungen an anderen Bandwuermen. IV.
Mitteilung. Ueber einige Diesing’sche Originale und
verwandie Formen. Stizungsber, Acad. Wiss. Wien
Math, Naturwiss. KL Abt. I, 138, 145-166.
ScHMIDT, G, D. (1986) Handbook of Tapeworm Identifi-
cation, CRC Press, Inc,, Boca Raton, Florida. 675 p.
Susyaprabua, C. K. (1955) Cestode parasites of fishes
of Madras Coast. Ind. J. Helminthol. 7, 41-132.
YAMAGUTI, S. (1934) Studies on the helminth fauna of
Japan. Part 4, Cestodes of fishes. Jop. J. Zool. 6, 1-112.
(1952) Studies on the Helminth Fauna of Japan,
Hak 49. Cestodes of Fishes, Il. Acta Med. Okayama
, 1-76,
(1959) Systema Helminthum. Vol. 1. The Cestodes
of vertebrates Interscience Publ, Inc., New York, 860 pp.
HORNELLZELLA MACROPORA (SHIPLEY & HORNELL, 1906) COMB.
NOV. (CESTODA: TRYPANORHYNCHA) FROM AUSTRALIAN
ELASMOBRANCH FISHES AND A RE-ASSESSMENT OF THE FAMILY
HORNELLIELLIDAE
BY R. A. CAMPBELL* & I. BEVERIDGEt
Summary
Hornelliella macropora (Shipley & Hornell, 1906) comb. nov. is redescribed from the
elasmobranches Stegostoma fasciatum and Chiloscyllium punctatum from northern Queensland.
H. annandalei (Hornell, 1912) is considered a synonym of H. macropora. The armature is described
for the first time and consists of a unique poeciloacanthous type in which large, alternating pairs of
hollow hooks form a double chainette on the external surface of the tentacle. The presence of an
hermaphroditic vesicle is confirmed and illustrations are provided. The family Hornelliellidae
Yamaguti, 1954 is considered justified, based on a combination of the unique features of the
armature and the genitalia, and is re-defined.
KEY WORDS: Cestoda, Trypanorhyncha, Hornelliellidae, Hornelliella, elasmobranchs.
HORNELLIELLA MACROPORA (SHIPLEY & HORNELL, 1906) COMB. NOV,
{CESTODA: TRYPANORHYNCHA) FROM AUSTRALIAN ELASMOBRANCH FISHES
AND A RE-ASSESSMENT OF THE FAMILY HORNELLIELLIDAE
by R. A. CAMPRELL* & I, BEVERIDGEt
Summary
CaMPBeELL, R. A, & Bryerines, | (1987) Hornelliella muceopora (Shipley & Hornell, (906) comb. nov;
(Cestodd: Trypanorhyncha) from Australian clasmobranch fishes and a re-assessment of the family
Hornelitellidae. Troms. R. Soc. S, Aust. 111(4), 195-200, 30 November, 1987,
_ Hornelhielia macropora (Shipley & Hornell, 1906) comb. nov. is redescribed from the elasmobranchs
Stegostoma fasciatyum and Chiloscy/liam punctatum from northern Queensland, H. annendalei (Hornell,
1912) is considered a synonym of A. macrapora, The armature is described for the first time and consists
of 4 unique poeciloacanthous type in which large, allernating pairs of hollow hooks forny a double chainetie
on the external suttace of the tentacle. The presence of an hermaphroditic vesicle is confirmed and illustrations
are provided, The family Hornelliellidac Yamaguti, 1954 is considered justified, based on a combination
of the unique features of the armature and the genitalia, and is re-defined,
Key Worpbs; Cestoda, Trypanorhyncha, Hornellietlidae, Hornelliclla, clasmabranchs.
Introduction
Yamaguti (1954) erected the genus Hornelliélia
and the family Hornelliellidae for a single species
of trypanorhynch cestode, Tefrarhynchus annart-
delet Hornell, 1912 described from a shark, Stego-
stoma fasclatum (Hermann, 1783) (syn, §, tigrinunt
(Cimelin)), from the Bay of Bengal. The new family
was distinguished primarily by the presence of a
unique structure within the reproductive system,
termed an hermaphroditic vesicle, which was not
illustrated, and was supported by several minor
characters including the distribution of testes and
vitellaria. In trypanorhynchan 4ystematics, the
tentacular armature is of prime importance (Dollfus
1942), but neither Hornell (1912) nor Yamaguti
(1954) described the armature in sufficient detail
to determine the taxonomic position of the family,
In addition, Southwell (1929) synonymised 7:
ennandatlie with Tentactilaria macropora (Shipley
& Wornell, 1906), a fact which Yamaguti (1954)
either overlooked or ignored. Cestodes of this
family have not been reported previously from the
Australian region, however, specimens have recently
heen collected from Sregostoma fascidtum and
Chiloseyllitim punctatum off the coast of
Queensland, In this paper the species is described
in Full for the first time, the taxonomic position of
the family Hornelliellidae reassessed, and the family
redefined,
* Department of Bivlogy, Southeastern Massachusetis
University, North Dartmouth, Massachusetts, 02747,
USA,
t Central Veterinary Laboratory, South Australian
Depuriment of Agriculture, Instilite af Medical and
Veterinary Science, Frome Road, Adelaide, S, Aust,
Materials and Methods
Cestodes from the spiral valves of sharks were
fixed in 10% formalin, and were stained with
celestine blue, dehydrated in ethanol, cleared in
clove oi! and mounted in balsam. ‘Tentacles were
dissected free and cleared in glycerol. Measurements
ate given in micrometres, unless otherwise stated,
as the range of 10 measurements followed, in paren-
theses, by the mean.
Terminology for trypanorhynch morphology
follows Dollfus (1942), Specimens have been
deposited in the Australian Helminth Collection
(AHC) of the South Australian Museum, Adelaide
Southweil’s specimens of Téntacularia macropora
were borrowed from the British Muscum (Natural
History), London (BMNH).
Hornelliella macropora (Shipley & Hornell, 1906)
comb. nov,
FIGS I-11
Tetrarhynechus trnandefe! Hornell, 1912
Aornellieta anancdie’ (Hornell, 1912) Yamaguti,
1954
Tetrarhynchus macroporus Shipley & Hornell,
1906
Tentacularia macropora (Shipley & ttornell,
1906) Southwell, 1929
Descriplion; Cestodes up to 76 mm long, with up
to 63 scements in gravid strobilae. Sculex
acraspedote, 7.25-11.2 (8.62) mm long, maximum
width at bulbs 0,85-1.02 (0.94) mm. Two bothridia.
1.7-2.6 (2.09) mim by 1,36, 1.38 mm, eval or slightly
dumb-bell shaped, notched posteriorly, rim broad,
median ridge within each bothridial cavity. Pars
196 Kk. A. CAMPBELL & I. BEVERIDGE
rr
ata,
SS
bah
grrrtrrs
nea cape :
1 a4 ‘
es)
8
=
Er}
a
fe oH)
Figs 1-6, Hornellietla muecrepora (Shipley & Hornell) from Sfegostomu Jusciaium., 1, scolex; 2, mature segment; 3,
genital system; 4, venital atrium with partial eversion of the hermaphroditic duct; 5, genital atrium with partial
eversion of the hermaphrodilic vesicle; 6, genital pore, lateral view showing lappéts anterior and posterior to pore.
Scale lines: Figs 1, 2: 1.0 mm; Figs 3-6: 0.1 mm. Legend: c, cirtus: ga, genital atrium; hd, hermaphroditic duct;
hy, hermaphroditic vesicle; isv, internal seminal vesicle; |, lappet; v, vagina.
CESTODE bAMILY HURNELLIELLIDAE 197
vaginalis 3,35-3.95 (3,82) mm, shealhs coiled, Bulbs
12 to J5 times longer than wide, 3,57-5.72
44.74) oun by U.28-0.38 (31) mm to diamerer:
prebulbar organs absent; retractor muscle attached
at anterior end of bulb: pars postbulbosa absent.
Soolen ol sharply delineated from strobila, merging
Inky neck.
Amiacure: pocciloacanthous, with a chainette of
iwo tuws of large, paired hooks on middle of
external face in metabasal resion; principal rows
alternate, forming ascending half spirals beginning
én Internal face: distinctive basal armature present;
no basal swelling; hooks hollow. Tentacle diamerer
130-170 (160). Basal armature: about cight
ascending balf spiral rows of small hooks on cach
side Of external surface; rows alteriate, Median,
teook-[ree area present on internal surface of base,
tapering toward metabasal region. Hooks !(t') of
proximal four rows of basal armature large, falci-
form with broad transverse bases; hooks I(I') of
basal rows 2-3 largest, 150-216 (184) long, base
44-90 (62); hooks 2 (2') also barge, recurved;
remaining hooks small, spiniform, opposing rows
meet 10 form inverted-V formations on external
face; spiniform hooks.of rows four-six larger than
tore distal or proximal rows. Basal armature
merges gradually into metahasal armalure. Meta-
basal armature: ascending tows with seven hooks
per half spiral. Hooks 1(1") widely separated, rose-
thorn shaped, length 126-160 (149), base 76-102
(93), smaller near base of tentacle; hooks 2(2°) large,
falcilormi, length 110-142 (13), base SO-74 (47);
hooks 33") slender, falciform, length 130-150 (136),
base 32-50 (40); hooks 4(4°%) slender, falciforn,
length 110-130 (124), base 20-30 (24). Hooks 5(5°)-
7(7') spiniform, forming triad, offset at oblique
angle and slightly posterior to hooks 4(4"; hooks
5(5'), length 88-10 (93), base 24-32 (26); hooks
6(6°), 82-110 (94) long, base 18-26 (24) hooks 7(7'),
1D4-120) (112) long, base 18-24 (21). Hooks 88) and
9(5°) widely separated from hooks 7(7') forming
ilternate pairs in chamerte, each pair opposite a
single principal row: Hooks 8(8') and 9(9"} spini-
forin, similar to hoerks 5(6')-7(7') of principal rows;
hugoks 88"), 72-112 (99) long, base 20-26 (23),
hooks 9(9°) smaller, 68-90 (73) long, base 14-28
(22).
Segments appear |.0-1,5 mm posterior to scotex,
initially segments very narrow, beconving elongated,
craxpedate; mature segments. 1,76-2.57 (2.10) mm
by 0,89-1.41 (1.18) mm. Testes numerous, 50-6! (50)
wi diameter, arranged in layers, occupying entire
medulla anterior to hermaphroditic sac, traversing
dorsal osmoregulatoty canals but not ventral pair
of osmoregulatory cattals; testes absent posterior
to hermaphrodithe sac, Vas deferens greally coiled,
distetided, occupying much of available space
between ovary and proximal pole of hermaphroditic
Sax, penctrating sac from anterior cde Hermaphro-
ditic sac, 520-700 (626) hy 250-390 (310), pyriform,
thick-walled, proximal pole deviated anteriorly;
contains crescentic internal seminal vesicle and
eoiled cirtus armed with tiny, aciculate spines.
Cirrus joins thick-walled, comugated, hermaphro-
ditic vesicle armed with prominen sagittate spines.
Hermsphroditic vesicle joins genital atrium through
short, muscular, cversible hernmiaphraditic canal,
Genital pore marginal, irregularly alrernare; genital
atrium deep, sitiated in posterior \4 of segment,
68-71% of segment length from anterior margin,
surrounded ameriorly and posteriorly by two large,
fleshy semi-lunar lappets, Ovary tetralobed in cross-
section, set forward from posterior margin of
segment, lobcs subequal, 130-250 (176) by 210. 290
(230), Mehilis' gland large, about 280 in diameter,
poslovarian, Vagina thick-walled, coiled, ascencting
from Mehlis" gland on poral side of vas deferens
fo penetrate posterior wall of hermaphrodiric sac
and join hermaphroditic vesicte. Seminal rooeptable
absent. Uterus simple, median, walls irregularly
lobulate, terminating near anterior extremity of
seument. Preformed uterine pore almost mid-way
between genital pute and anterior margin of
segment, Vitellaria follicular, crcunicortical, (ing
Pastovarian space, surrouriding all internal organs
except most anterior testes, Eeps ovaid, pale in
colour, Longntudinal osmoregulatory canals paired,
ventral canals with transverse commissure at
posleriur margin of segment: dorsal canal medial
to ventral canals.
Materiat examined: 11 specimens [rom spiral valve of
Stegosionwy fasciaiurnr (Hermann 1743), Balgel, Olu,
L4.ix.1985S, coll. BOG. Robertson ($2771); L specimen trom
Chiloseyllium puneraram Mucilot & Henle, (841, Balgal,
Old, 17,i4,79K5, voll. B. G. Robertson ($2772).
Discussion
Southwell (1929) synonymised Télrarsynchus
macroporus and T. athriundate? under the combina-
tion Tertecwluria macropora, having examined a
range of specimens from different host species,
including the type host of 7: emandalet, Shipley
& Hornell’s description (1906) of Ferrarkvichis
miaccropariks was based on specimens fron Fintan
tura uarnak (Forsskal, 1775) (Syn, Tregon uemek),
The whereabouts of their specimens is unknowu,
and no specimens have since been recovered from
thts hast, Their description of the species is brief,
und as Southwell (1929) pointed out, differs. little
from thaLof 7 annanda/e:, Southwell (1925) con-
yidered jhe difference belween (he two species,
namely the presence or absence of a longirudiral
R. A. CAMPBELL & J. BEVERIDGE
198
Nee
=
Se
<——
ss
{
2
internal surface, metabasal region;
face, basal region; 11, profiles
internal sur
?
,
figs 7-10 same. scale.
’
nelliella macropora (Shipley & Hornell). 7
hothridial surface, basal region; 10
E
E
ies
>
nae 2 |
sf
SEs
O23
vos
S329
3
sw
= 4
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ie
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te
CESTODE FAMILY HORNED TTELLIDAE 199
division uf the bothridium, to be due to intraspecific
variation, and noted thar contracted specimens are
mote likely to have @ medium sub-division of the
bothvidia, Southwell’s (1929) cowclusiuns are
considered justified since if specimens are examined
unstamed using incident light, there is a prominent,
media longitudinal ridge in each bothridium,
which becomes virtually invisible when the speci-
mens are cleared, Hornell (1992) also stated that the
length of “proboscis hooks" differed between the
Iwo specics, though no measurements were given
in etthec of the original descriptions Marked
differences occur in hook size and shape ona single
tentacle, and hence such comparisons are useless
if the posilion of the hooks being compared is not
clearly specified.
Hornell’s (1912) specimens of f! annuidalet were
taken froin Stegastenia fasciatwnr (Hermann, 1783)
(syn, 3, ngrinwen (Gmelin, 1789}), as were those of
Yamaguti 1/934) and the new specimens described
herein, The material examined by Southwell (1929)
included specimens from S. fasciatum rom Sri
Lanka (BMNH 1977 11.14.46) as well as speamens
fron) Guteocerdo cuvier (Peron & Le Sueur, 1822)
isyn, Gi wretions (Faber, 1829) and G. tigrinus
Mueller & Heale, (839) from Sri Lanka (BMNH
1977.LL.14 32-35) and specimens from a host
identified on the label simply as 7rygen sp. (BMNH
L977.01,14.9 aid 1977.11, 14, 21-24) bur identified in
the text of his paper as Dasybenes sp. All specimens
clearly belong to a single species, with a wide host
range, We therefore support Southiwell'’s (1929)
conclusion that 7) niacreporus and 7) annandalei
are conspecific though material trom Himanture
uarnak is needed to confirm this view. Yamaguti
(1954) appear to have overlooked this synonymy
ht his redescription of T; wnaendale’, and hence the
correct combination for his new genus becomes
Hornetliella macropera (Shipley & Hornell, 1906)
comb. nov. (syn. 4, annandalei (Hornell, 1912)
Yamaguri, 1954).
Hornejlielia palasoorani Zaidi & Khan, 1976 is
the only other nominal member of the genus, The
spevies was said lo.differ from H. micropora only
in the size of the cirrus sac (Zaidi & Khan 1976).
However, the description is very poor arid itis clear
fron the drawings thal it does not belong to the
gens Hornellielia, is systematic position cannot
be determined and it is considered fneertae sedis,
the description given herein continms most of
Yamaeuti's (1954) observations, except for the single
seminal vesicle which appears, from his description,
to be external ro the virrus sae bue whivh in faet,
is inlermal (see Fig. 3).
The artnuture, desertbed Here for the First time,
presents several unique featuses. The large Falcifori
hooks of the basal armature are similur w species
of Gramoriyachus Rudolphi, [819 and Malirole
(Pollfus, 1935} Also, the external surface of the
metabasal armature bears what we have described
as a double chainette of hooks which are similar
in form (o those of the principal rows, The hooks
ol the chainette are well separated from and
opposite to the alternating principal rows as Occurs
in genera such as Lacistorhynchus Pintnes, 1913 and
Caliiterraraynchus Pintner, (934. All of che hooks
are hollow and are not accompanied by satellite
hooks or by intercalating rows of hooks Although
a4 poecifoacanihous type of hook atrangement
fiiting the orderly arrangement found in a chainette,
it could also be interpreted as a “hand” of hooks
such as is found on the external surface of the
tentacles of species Molola and Grilloria (see
Dolifis 1942). However, such bands of hooks in
poectloacanth trypanorhynchs {reguently shove little
or no orderly linear arrangement of their elcmenrs
and the haoks may vary both in size and in form,
We therefore prefer ts consider the hook
arrangemenc seen in Hornelfiglig as a modified
chainette.
A double chainetie with hooks in a tandem
position, acours in Lacistarkynchas and Eulaciste-
rhynchus Subhapradha, 1957 (Lacistorhynchidac),
in Callitetrarhynchus (Dasyrhynchidaes, Gymno-
fhunchus (Gymnorhynchidae) and some species of
Dusprkvnichus Pintner, 1928 {Dasyrhynchidae).
Simple chainettes with hooks in a single row oceur
in FYaviceps Cuvier, 1817 (Dasyrhynchidae), Hely-
siockyachus Pinner, 1913 (Pierobothnidae}, Afixo-
digma Dailey & Vogelbein, 1982 (Mixodigmatidae)
and in same species of Dasyrhiyrcius, No genus
described to date has a chainette composed of pairs
of hooks in tandem, and the structure seen in
Hernellietia is unique among the Trypanorhyncha_
Yamaguti (1954) erected a new Family and genus
for the species hased primarily on the presence of
a unigue, hermaphroditic vesicle, which be chd not
iNustrate. Yamuguti's description is correct, and
both cirrus and vagina join a large, thick-walled
vesicle armed with sagiltate spines The vesicle leads,
via a highly muscular hermaphroditic duct, to the
geniial atrium, A fully everted circus was not
observed but the hermaphroditic duct appears to
be éyersible and the vesicle is capahle of being
partially protcuded through the duct, with the
characteristic sagittate spines being visible on the
external surface (Fig, 5) Althougl) the vesicle
appears to be unique, a union of male and female
ducts within the “cirrus sac” |=hermaphrodiue
pouch of Yamaeguti 41954)) oeeurs in Lectsre-
rAynchus (see Dollfus 1942) hut in the latier case
there is no armed vesicle, merely a simple
hermaphroditic duct referred to as the “gentlal
atrium by Doilfus (1942) because Ihe “arriwin" by
200 &. A. CAMPBELL & |. BEVERIDGE
everted first, followed by the cirrus. A similar
mechanism of evagination may operate in A.
niacropora. We consider the terminology used by
Yamaguti (1954) of hermaphroditic duct. and herma-
phroditic vesicle preferable to that used by Dollfus
(1942) for Lacistorhynchus tenuis.
Yamaguti (1954) erected the family Hornellielli-
dae based on the preserice of an hermaphroditic
vesicle, testes anterior to the “cirrus sac”, presence
of paired lappets around the genital atrium, a series
of muscular rings in the tentacle sheaths and
circumcortical vitelline follicles which formed a
band posterior to the ovary. Of these characters,
the muscular rings of the tentacle shcaths were not
visible in our specimens, and we doubt whether such
precise distribution of the testes and vitellaria will
prove to be family characters. However, the
armature is unique in that each pair of hooks in
the chainette is contributed by a single, principal
row, in alternating fashion. Furthermore, the hooks
of the chainette are similar in shape to those of the
principal rows instead of being markedly different,
On the basis of its armature and the herma-
phroditic vesicle we consider that the Hornelliellidac
is valid and that it is most closely related to the
poeciloacanthous families Lacistorhynchidae
Guiart, 1927 and Gymnorhynchidae Dollfus, 1935.
A redefinition is given below,
Hornelliellidae Yamaguti, 1954. Hornelliella
Yamaguti, 1954, Scolex elongate, acraspedote; two
oval bothridia each with median longitudinal ridge.
Metabasal armature poeciloacanthous, chainette of
two rows of large paired hooks; intercalary hooks
absent; distinctive basal armature present; no basal
swelling. Tentacle sheaths coiled; prebulbar organ
absent; retractor muscle originates in anterior third
of bulb, Pars postbulbosa absent. Genital pores
marginal, irregularly alternate. Testes numerous,
scattered, anterior to genital pore; internal seminal
vesicle present; male and female ducts unite to form
armed hermaphroditic vesicle; genital pore sur-
rounded by lappets. Vitellaria follicular, circum-
cortical, pre and post-ovanian. Uterus simple,
median, tubular; preformed uterine pore present.
Parasites of sharks and rays.
Acknowledgments
Thanks are due to B. G. Robertson for collecting
the specimens, and to Dr D, Gibson (BMNHB) for
lending specimens.
This project was supported financially by the
Australian Biological Resources Survey.
References
Dor.rus, R. P. (1942) Etudes critiques sur les Tétra-
rhynques du Muséum de Paris. Arch. mus. nat, Hist.
nat, Paris, 6° sér. 19, 1-466.
Hornet, J. (1912) New cestodes from Indian fishes. Rec.
Ind, Mus. 7, 198-204, plates IX, X.
SOUTHWELL, T. (1929) A monograph of cestodes of the
order Trypanorhyncha from Ceylon and India, Part 1,
Spolia Zeyl, V5, 169-312.
Yamacuti, 5. (1954) Parasitic worms mainly from
Celebes, Part 6. Cestodes of fishes, Acta Med. Okavama
8, 353-374, plates I, 11.
ZarpL, D, A, & KHAN, D. (1976) Cestodes. of fishes from
Pakistan. Biologia 22(2), 157-179.
NEW RECORDS OF UPEROLEZA (ANURA: LEPTODACTYLIDAE) FROM
WESTERN AUSTRALIA WITH SUPPLEMENTARY OSTEOLOGICAL
DATA ON UPEROLEIA MICROMELES
Summary
BRIEF COMMUNICATION
NEW RECORDS OF UPEROLEIA (ANURA: LEPTODACTYLIDAE) FROM WESTERN
AUSTRALIA WITH SUPPLEMENTARY OSTEOLOGICAL DATA ON UPEROLEIA
MICROMELES
Upereleia Gray, 1842 comprises 23 speeies! of which
ten o¢cur in Western Australia: two from the Pilbara and
eight from the Kimberley Division, Recently sianificant
range extensions al two species inio W.A. have been
discovered.
Uperoleia trachyderma Tier, Davies & Martin, 1981
On 13..1987 we collected LU! Irachyderma during heavy
thunderstorms on the irrigation flats of the Ord River
Scheme at Kununurra (WAM R96781, 8.5 kin NNE
Kununurra, at the junction of Ivanhoe and Research
Stations Rds). ‘his represents an addition to the fauna
ot WA,
The species was collected on self mulching grey cracking
clay.* U. trachyderma is the only member of the genus
ta be associated with a particular soil type, and all
previously collected specimens have this fidelity,
Self-mulching grey cracking clays are distributed
disjunctly throughenut W.A., the N-T. and Qld;* those on
the Barkly, Tableland are considered to have evolved in
stv.? Distribution along current drainage patterns would
explain the presence.of the species‘on the Georyina River
and at Lawn Hill in Qld! but the occurrence of the
Species at Newry Station in the far northwest of the
NJ) and at Kununurra remains an enigma,
Two other vertebrates havé comparable disjunct
disteybutions on grey cracking clays: the monitor baranis
spercert’ and the planigale Planigale ingrami\®* while
two specics of bird, the flock pigeon Histrigphups
Aisteionice and the red-chested quail Turnix pyrrothorax,
have theic centres of abundance on these soils. Perhaps
jhe question of the origins of the Barkly Tableland soils
sliould be examined in the hight of the endemicity of the
associared fauna.
An additional species of Uperoleia has been recorded
in W.A.* A specimen of 1) mivromeles was collected by
A. A, Burbidge and P. JS. Fuller on 25.iv.1979, 1 km S
Stafford; Bore (22°22'S, 127°24'E) (WAM R64073),
Previously known only fron) {he Tanami Sanctuary, NT.
the species is unusual in 4 number of niarphological and
ostcologica!l features. Re-examination of the paratype
alizarin has enabled the clarification of same osteological
features not included in the original description! Jiere
we redefine the species and report these fealures.
Uperoleia micromeles Tyler, Davies & Martin, 198)
Definition: A relatively large species (@ 27 mm, 9 9
24-31] mm 5-V) of gross habitus; maxillary teeth present
ap vestiges; narrowly exposcd frontoparietal tontanctie;
short, fattened unweébbed toes; internarial span greater
than cye to naris distance; carpus of six elements; itial crest
present. Advertisement call not known.
Osteology: Uiunr with small dorsal crest (Fig. 1). Dorsal
prominence very large, mesa-shaped. Dorsal promberance
laleral; pubis caleitied,
Carpus of six elements. ©. radiale larger than ©. ulnare.
These elements articulate proximally with G. radiautna,
with each other on their proximomedial border, with large
transversely elongated O. centrale postaxiale distally. 0.
radiale articulates laterally with O. centralé preaxiale.
ae SP,
Ao — mR | 7
Fig. 1 Lateral view of the pelvis of Upercleia micrameles
(Paratype):
O, centrale pastaxiale articulates distally with bases of
O. metacarpi UI, IV and V; small flange extends
proximally from. lateropraximal comer (Fig. 2). O. centrale
prexiale articulates laterally with O. radiale, distally with
©. centrale postaxiale and carpal elements of O. distale
carpale 2.and 3, laterally with basal prepoltical element.
Three distal tarsal clements. Lateral element largest, lies
at base of O, metatarsus If], extending laterally to
articulate with medioproximal side of base of ©.
metalarsus 1, medially to base of O. metatarsus 1]. Medial
element lies at base of Q. metatarsus I, articulates with
QO, centrale prehallucis, Distal prehallical element long,
elliptic, extending for approximately 2/3 the length of ©,
metatarsus | (Fig. 2),
1, |
4
Bal
f
Biz. 241A) Palmar view of the carpus and (B) plantar view
of the tarsus of Uperaleia micromeles (Paratype).
202
Comparison with other species: Uperoleia micromeles is
a functionally edentate species (the teeth are vestigial) with
a poorly exposed frontoparietal fontanelle. This species
is unique amongst Uperoleia in a number of features:
presence of vestigial teeth; a broad snout (E-N/ IN
0.83-0.90!!); anteromedial extensions to the nasals; and
the presence of a moderately developed ilial crest.
We thank Michael J. Tyler for reading the manuscript,
John and Judy Caratti for hospitality in Kununurra, and
Chris Done and the W.A. Department of Conservation
and Land Management for field assistance, This work was
supported by research funds from the Departments of
Zoology, Universities of Adelaide and Melbourne.
\Davies, M., McDonald, K. R. & Corben, C. (1986)
Proc. R. Soe, Vict. 98, 147-188.
2Northcote, K. H., Hubble, G. D., Isbell, R. F,
Thompson, C. H. & Bettenay, E, (1975) “A description
of Australian Soils,” (CSTRO, Melbourne).
3Davies, M. (1987) Taxonomy and systematics of the
genus Uperoleia Gray, 1842 (Anura: Leptodactylidae).
Ph.D. thesis, University of Adelaide. Unpubl.
4Northcote, K. H. Pers. comm.
Tyler, M. J., Watson, G. F, & Davies, M. (1983) Trans.
R. Soc. S. Aust, 107(4), 243-245.
6Tyler, M. J. & Davies, M. (1986) “Frogs of the Northern
Territory.” (Conservation Commission of the N-T., Alice
Springs).
7Pengilley, R. (1981) Aust. J. Herp. 1, 23-26.
8Parker, S. A. (1973) Rec. S. Aust. Mus. 16(11), 1-57.
°Andrew, D. L. & Setile, G. A. (1982) pp. 311-24 Jn M.
Archer (Ed.) “Carnivorous Marsupials.” (R. Zool. Soc.
N,SW., Sydney).
10Parker, S, A, Pers. comm.
'lTyler, M. J., Davies, M. & Martin, A. A. (1981) Aus.
J. Zool. Suppl. Ser. 79, 1-64.
MARGARET DAVIES, Department of Zoology, University of Adelaide, G.P.O. Box 498, Adelaide, S. Aust. 5001,
GRAEME F. WATSON, Department of Zoology, University of Melbourne, Parkville, Vic. 3052 and CHRISTOPHER
A. MILLER, Department of Zoology, University of Adelaide.
AUSTRALIAN PLANT NEMATODES: LONGIDORUS MICOLETZKY, 1922
AND PARA LONGIDORUS SIDDIQI, HOOPER & KHAN, 1963
(NEMATODA: DORYLAIMIDA)
Summary
BRIEF COMMUNICATION
AUSTRALIAN PLANT NEMATODES: LONG/DORUS MICOLETZKY, 1922 AND
PARALONGIDORUS SIDDIQI, HOOPER & KHAN, 1963
(NEMATODA: DORYLAIMIDA)
Ina comtinuing study of the distribution and laxonomy
of plant parasitic nematodes associated with naturally
occurring native vegetation, spesies of Curu/onagidoeus and
Longidorus were identified. The distribution of both these
genera in Alistratia is poorly known. All records here are
from areas Gt natural vegetation, many of which are
National Parks (NP), Conservation Parks (CP) and Stale
Forests (SF) (excluding Plows redsata and other planted
forests). Nematode specimens are held in rhe Waite Insri-
tuic Nematode Collection (WINC). Measurements are
fram specimens mounted in glyceral.
The genus Lonaiderus has been recorded from three
locuboms around Sydney (NS SAV)" and from Charleville,
Qld? bur npne js identified to species, Soil samples
collecied fram four localities in South Australia contained
speviniens Of Lomgiderus faniwha Clack, 1963, a New
Zealand spovits, not previously recorded in Australia. The
localities are all on the Fleuricu Peninsula, sourh of
Adelaide, in Eucolyivus L'Hér,, forest at Mt Compass,
Kyeerna CP and Spring Mount CP, and wn area of mallee
ut Myponga CP. Apart from a more posterior guiding ring
(52-37 pm from anterior end), South Australian speeinens
arc similar in most respects lo the type population. Small
numbers of males were cotfected from each locality except
Spring Mount.
Three species of Purglongidorus have been recorded in
Australia. Two of these, P evcalypti Pisher, 1964 and
# saechar! Siddiqi, Hooper & Khan, (963 have been
idemlified by the aulhory, from bus{land soils, A euca/ypii
has only previously been necarded Irom the type locality
(S. Aust.)) and three localities in N.SW.! Specimens of
P evedlypt) have been Wentified from mare than 35
dilferemt Ineulities, in all States of mainland Australia.
Most of the localities are Eucalypius woodland or mallee.
rarely torest, none Irom rainforest. Soil samples have betn
collected from 24 sites, within 14 separate greas of rain-
forest (totalling (76 soil samples) in exstern Qld, and
NSW, No species of Paraloneldorus have been recovered
from Uiese soils.
fomales of Hardiongidaras eucalypi{ were swudied, and
the marphometnes of specimens from eight lovalines are
elven in Table b, The specimens are similar to the vriginal
description, with minor differences in tail length and
shane. The papillac near the amphid apertures, a peculiar-
Hy of (Mis species, ace clearly visible in mest specimens.
There ale seme clifYerences between measurements of
specimens fram differing lavalities, The $, Aust, specimens
ate similar to those from the type lecality, The Qld
ypeciens are shofter (han those from S. Aust. and also
have a shyrter styler and a more unleriorly situated guiding
ring. Some specimens trom Marne Valley 1S. Aust.) and
Mt Topper (N.SM.) have a slightly more pusteriorly situ-
ated vulya. One female specimen, identified tram
Dengarra (WAL) Las an untoually short stylet (127 pm)
but is similar in other respects to f% excalypti. The
messurements given here considerably extend the range
for This species,
Localities from which # eucalppti were collected are
48 follows:
S. Aust: Guileys to the vast and west of Northern Minders
Ranges, in association with Evcalypius cumuldulensis
Dehnh; mid-north, associated with Liaulypius woodland
and mallee; frequeul on sandy sous associated with
Fucalypius woodland and mallee, including: Saudy Creek
CP (neat Gawler); Ferrics McDonald €P (near Murray
Bridge); Marne Valley CP (near Swan Reach): Aldinga
Serb (Adelaide cowl).
Vie, Sunraysia district, associated with Eucalypius
comuldilensis oy mallee; Eucalyptus woodland at Victoria
Valley, Grampian Mountains,
NSW: Sunraysia district, north-castern State Terests
including M1 Topper SF; Sacks Creck SFS Nundle SF,
Unumyar SP; Beaury SP,
Old: Eveaiypies woodland west of Mt Garnett (north
Old: brigalow asseciatian, Thangool (near Biloclay:
Cosuerina ditterefis Salish, fuulside rainforest area)
Eungella NP near Mackay); malice and mutpa,
Charleville
NT Bucwlvpius woodland with tall perential arasses,
Katherine.
HAs Buedyeres calaphylla R.Br west af Narrham,
E& erythrocorys UT. Muell., south of Dongarra (near
Geraldton),
The male of # evcalypr/ has or be desecibed, One male
has been collected from S. Aust. and qne from W.A., which
are described here.
Measwrements (5. Aaist. male, n = ib - 3 S8 mm, a
~ M27 p0 = 8.2; ¢ — 128; odonmtostyle ~ 106.4 pms
adoniophore = 49,5 pms total stylet = 155.9 am; guiding
ring 43.4 ym [rom antetior end; T = 63.4; spicules
= 43.4 pm (tivasured wlong curvature): lateral guiding
icces 14,7 pms tail = 43.5 pm,
(WA. male on = Lb = 539mm - 12L%¢ - 223
ec = 135: odoninstyle - IDL2 ym; oadantophore
= 46.8 pin; total srylee = 144.0 pin; guiding ring 34,9 am
from anterior end; T — 48.0; spicules 56.3 am), lateral
guiding pieces LI1 pm; tail ~ 44.03 pm.
Neserlotiag: Body ventrally Curyed whien relaxed (Fig. LA),
more strungly curved in posterior region than fernales,
Anterior region similar ta temales, with offset lips region
(Fig. (BC), papillae near amnphid apertures and stirrup
shape! amphids. Odontostyle shorter rhan females fram
type populaiion. Hemizonicd 3.8-6,8 por long, Cinnas!
rather obsetire. Spicules arcuate, lateral guiding pieyes
slightly cutved. In the S. Aust. specimen, supplements
consisting of ay adanal pair, unequal distances from the
nus, with a venrra-median scri¢s of fourteen. If the W.A,
specimen there aré thirteen ventro-median. supplements,
The most anterior supplemeiy is $5.1-95,2 pm from anus
(mewured along curvature), Tail rounded-conoisl terminus
slighty bulbous in buth specimens (Fig. 1D, 2).
Localities: (8. Aust.) Collected fram Aldinga Serub,
S. Aust, in December 198] by (he author Uhts site is law
‘pus JOWAIUe WOI] Aus BuIpins Jo aoueisip = “VST = Us
atwest (ir-2€) (ET) (Zrt-971) fhTT-301) (Gr-Br) (E66) (g-9P) (9¢I-SszD = (€8t-1z1) ('s-1°9) f CIN)
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asit = FOE 60 6 P81 €'S0T 6 iP gir 8°05 g6rl 9007 99°¢ 1 gnisg eauipry
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(uu) (ur) (un) (uu?) (ut) (mur)
ONIM ye a 2 eB A 1d aroydowwopg = ayAyso}wopO TBIAIS TI u Aye07}
‘sainyv20] auiu woif Wdsjeons sniopisuojeeg fO Sodjausoydsopy “| ATVI,
D
Fig. 1. Paralongidorus euculypti—male. A—relaxed body
posture; B, C—lip region; D—posterior region; E-tail.
Rar seales: A = 1 mm, B-E = 20 ym,
lying, sandy soil with Eucalyptus cumaldulensis and
numerous Isolepis nodosa (Rotb,) R.Br. (Knobby Club-
Rush) WINC: 176A,
(W.A.) Male and juveniles collected from dam reserve at
‘The Humps”, Hyden, W,A,, by P. Orr and P. Laidlaw
in September 1986. Caswarinad sp. dominant, with
associated Acacia sp, Eucalyptus sp. Santalum
acuminatum (R.Br) A.D.C. and tussock grasses, WINC
332A.
' McLeod, R. W. (1979) Science Bulletin 87, 48 pp. Dept
Ap. N.S.W-
> Khair, G. T. (1987) List of plant parasitic nematodes of
Australia (3rd Edn) Canberra, Dept of Primary Industry.
156 pp.
3 Fisher, J. M. (1964) Nematolopica 10, 464-470.
205
Paralongidorus sacchari, first described from north Qld
and India‘ and also recorded fram FijiS has not been
recorded elsewhere in Australia. Specimens have been
identified from two localities in S, Aust, and are
morphologically similar to the original description, but
differ in same measurements.
Black Rock CP, females (a = LO). L = 3.7-5.5 mm;
c = 100-141; co = 1.1-1,3; V = 47-52; odontostyle =
119-134 pm; odontophore = 36-60 um; total stylet =
162-19) pm; guiding ring 32-37 ym from anterior end:
tail — 30 44 pm.
Locality. Black Oak, Casuarina cristata ssp. pauper (P.
Muell, ex Miq) L. Johnson and Blue Bush, Maireana
sedifolia (F. Mucll), P, G, Wilson association at Black
Rock CP (near Orroroo) S. Aust.
Females and juveniles collected in 1974. WINC 59-63.
Martins Well, females (n = 3). I, = 3.7-4.8 mm; c¢ =
109-157; c = 1.0-1.2; V = 48-50; odontostyle = 115-130
um; odontophore = 38-62 gm; total stylet = 154-192 pm;
guiding ring 35-383 wm trom anterior end; tail = 30-35
pITL,
Locality: 118 ktm south-east of Blinman, towards Martins
Well (ouiside northern Flinders Ranges), from gully mar-
gin. Associated vegetation includes Eucalyptus
camaldulensis and EB, interfexta R. T. Baker, and Ptilpius
spp. Females and juveniles collected in 1973. WINC. 29
A, D.
Sincere thanks to people who collected soil, in particu-
lar; J. A. Simpson, Forestry Commission of N.SW., R.G.
Silcock, Dept of Primary Industries, Qld, M, H, Andrew,
CSIRO Division of Tropical Crops and Patures, N-T., J. H.
Warcup and P. McGee of the Waite Institute, P. Laidlaw
and P. Orr, Adelaide. Thanks also to the 8, Aust. and Qld
National Parks and Wildlife Services, $. Aust. Planning
Commission and the Forests Commission of Victoria for
co-operation with soil sampling in their reserves, ‘Technical
assistance by P, Fuhlbohm is gratefully acknowledged.
This work was partially funded by a grant from the Burcau
of Flora and Fauna,
4 Siddiqi, M. R,, Hooper, D. J. & Khan, BE. (1963)
Nematologica 9, 7-4.
> Orton Williams, K, J, (1980) Plant parasitic nematodes
of the Pacific, Tech. Report Vol. 8. 192 pp.
Commonwealth Inst. Helminthology, England,
FRANCOISL REAY, Department of Plant Patholovy, Waite Agricultural Research Institute (University of Adctaide),
Glen Osmond, 8. Aust. 5064.
YABBIES AT DALHOUSIE SPRINGS, NORTHERN SOUTH AUSTRALIA:
MORPHOLOGICAL EVIDENCE FOR LONG TERM ISOLATION
Summary
BRIEF COMMUNICATION
YABRIES AT DALHOUSIE SPRINGS, NORTHERN SOUTH AUSTRALIA:
MORPHOLOGICAL EVIDENCE FOR LONG TERM ISOLATION
This paper stems from a comprehensive morphological
analysis of the taxonomy of the @estrucor group of the
genus Cherax,'* Since the specimens from Dalhousie
Springs proved to be morphologically distinct from all
other specimens examined, they are given special
consideration here,
Dalhousie Springs are u series of over 60 active mound
springs distributed (throughout an area of about 60 km*4,
160 km NE of Oodnadatta in the far north of South
Australia? While located within the Lake Byre Drainage
division, they are isolated from the major waterways and
lakes of central Adstralia.
Morphological analyses included both bivariate
(analysis of covariance) and :multivariate (principal
components analysis) techniques? On the basis ot tity
metric and multistate characters, the specimens from
Dalhousic Springs are closest to C. destructor, which is
the only other species of Cherax ty be found in central
Australia.? Of these characters, the Dalhousie Springs
specimens could be distinguished sratisiigally from
C. destructor by iheir relatively broad arcola (the distance
between the branchincardiac groaves), short thorax, and
Shert vostuim, and to a lesser cxtent by relatively Jarge
chelae and broad rostrum.’ These characters are
iWusttared in Fig, 1,
The Dalhousie Springs spevimens are separated nrost
conspicuously frome. desiructor by their relatively broad
areola. The relationship between areola width and ocular-
carapace length for the Dathousle Springs specimens (a),
C. albidus (b) and C. destructor (c) is shown in Fig. 2.
The data for Dalhousie Springs specimens represent
collections from several mound Springs. These data were
tog-itansformed to linearize the allometric relationship of
areola width with ocular-varapace length, Analysis of
covariance indicated that the elevations of these regression
lines are significantly different (F497 =6)1.), p<.0,001),
A pesteriori comparison of the elevations using Tukey's
Tes!” indicated that the differences between each pair of
lines are statistically significant (p<0.05), with the
Dalhousic Springs specimens having the broadest aveolac
and C. destructor having the narrowest.
lgalhousie Springs are thought to have formed during
the lale Pliocene-early Pleistocene and Ure surrounding
atea hay been substantially dry sinve the late Tertiary.’
However, in modern times, flood waters from the creeks
near the springs would have provided a connection
between the springs and the Finke river system via spring
run-off during particularly wet years.’ Tt has been
suggested that these canditions may have occurred
nurnerous times in the past 1000 years and possibly more
Frequently during the Pleistocene wher the springs niay
have been ata ictatively lower elevation.’ In the hzht of
these suggestions, it is interesting that these populations
have maintained their morphological integrity, probably
in the face of repeated invasion by C destrucrar. One
explanation for this may be that rhe yabbies of Dalhousie
Springs may have developed pliysiplogical tolerance of the
high, relatively constant tumperiture to which they are
eaposed, Invading C. destructor may not be ahte to survive
+——
TGL
hig, 1, Cephalothoracic measurements distinguisling the
Dalhousie Springs yabbies from arher specimens of
C. deytruvior, TCL, total-carapace length; OCL, aculur-
carapace length; TL, thorax tength; RW, rostron. widit;
AW, areala width. Rostrum length was calculated as the
difference between TCL and OCL.
and reproduce under chese conditions. Ecological isolation
af populations hetween which there are no geopraphical
barrienm has been described tor some species of Garnmnartes
which inhwhit regions with differing salinities.*
Individuals from different populations of some “species”
were 50 specialized that they could not survive and
reproduce at any common salinity.
On the basis of zoogeographic evidence not connected
with Dalhousie Springs, | have previgusly postulated that
¢. destructor arose as a result of increased selection for
drought tolerance during the late Teruary-Pleistocene
phase of increasing aridity and that it arose from a
previously widespread “C. albidus-like!! ancestov,'? This
latter species is now restricted to southwestern Victoria
and (he extreme southeast of South Australia. The
similarity of the Dalhousie Springs specimens to
¢! alhidus, wlth respect 1o relative areola width is,
therefore, particularly siznificant. Mound springs also have
been sugyested as providing refuges: for aquatic fauna
LNUAW tant
26 Za 2.0 32 34 3.6 3.8
LATOCL mimi
Fig, 2. Lines of best Jit for the log-log regressions of areola
width on oculat-cvarapace length. a, Dalhousie Springs
in=75); h, C. albidus (n=526, <. C. destructor (n=900).
All regressions ate statistically significant (p<0.001).
The circles indicate the positions of individuals from
Dalhousie Springs.
dunng periods of wiereased aridity.? An association
between dreola width and the environment has been noted
for cambarid species, wide arcolae being largely restricted
to highly oxygenaled Jotic habitats and narrow areolar
being more common on crayfish inhabiting waters of lower
oxygen concentrations,” This association was related
to the fact that a lengthening and narrowing of the areola
will result in a concomitant enlargement in the gill
chamber? We wight therefore predict thar 2 drougit-
adupled spevies (ie. C. destructor) would have a relatively
narrow angola, ASC. des{reector populauans inhabiting
the cooler. wetter areas of central Victoria and
scatheasterm New Soulh Wales do nol show any
broadening of the areata, !* this relationship (if it applies
lo parastacids) ts not & proxinoal ore
Although the Spring water lemperatures are high,
dissolved oxygen levels are generally high ti saturated m
most springs, except in the immediate vicinily of the
vent,” The permanent water of the mound springs would,
therefore, buffer the crayfish fron: che effects of undity
and subsequent selecrian for drought tolerance, For further
interpretation of the pattern of morphological variation,
it would be interesting to Know wherher there has been
gene flow between the Dalliousie Springs crayfish and
typical C destructor. Future investigation using other
methods (c.g. immunological techniques} may help 1a
larity the relationships between the Dalhousie Springs
populations, CQ destructor and C albiadus-
Although these specimens are murplioloyically very
distinct from the other C destructor specitnens, their
taxonomic statusis far from clyar, The possability that their
distinetive morphology is environmerstally induced must
be considered, The Dalhousic mound springs represent
an exireme environment. Spring watet temperatures at the
source vary from 29°C ta 44°C, the hotter springs betrg
in the north. While summer water temperatures may
reach these levels in Gther parts of central Australia, che
Spring water (emperatires are constant throughout the
year it least at ihe source. These springs rarely Naw more
thad 1S km from their source due Ww the high evaporation
rate (2500 msn /annum), "1s the unusual morphology of
the Dathousi¢ Springs. specimens is environmentally
induced, some indication of morphological similarity with
other central Australian populations might be expected_
However, with respect to the chatacters which distinguish
the Spring specimens, yabbies from other nocthern Sauih
Australian, southern Northern Territory and southwest
Queensland localities are eloser to southeast Australian
specimens,!*
A further indication of the extreme environment at
Dalhousie Springs is the presence of berried females of
very small carapace lengths. | have examined berried
females of tolal carapace length (TCL) 23.4 inm (OCL
19.8 mim) and 24.3 mm (OCL 20.6 mm) fram this locality
Previous field studies have recorded miaimum total
carapace Jengths (TCLs) for bertied females of 31-
45 mro," although berried females of total carapace
length o& small as 26 mm have been revorded from Lake
Eucumbene in southeastern N.SW.'" The Dalhousie
Springs specimens therefore represent the smallest. berried
fernales of “Ct desrructor™ yet recorded. The size at which
crusiaacans reach sexual maturity in the field has been
Sugpested to be a guuge of (he influence of the
environment on the performance of the papulation,'®
A further complication is the possibility of recent
introductions. Yabbies were first recorded from the Springs
in May 1984 when they were collected using wire-mesh
box raps for fst!" However, bo Cherak were collected
on previous visitzin July/August. 1968, November, 1969,
and May, 1974 using the same raps in the same area.
When collecting vabbies, hawever, itis nor unusual to have
markedly different trapping success at different times. It
has also been nated that locals inthe mid-north of South
Australia, at least as far north as Godnadatta, have
introduced yabbies into their dams.!? Given that the
morphulogy of the Dalhousie Springs crayfish is so distinet
from all other C desiructer populations examined,'* it
is most untikely that they are the resule of recent
introductions by man
While the frequency and extent of past aquatic connec-
tions between ihe Dalhousie Springs and surrounding
drainage systems are not known, conditions Tavouring
speciation of other taxa have existed since there are two
endemic fish species, the Dalhousie catfish (Neosilurus
spt und the Dathousle hardyhead (Careracephulay
dalhousensis) whict cannol survive al waler lemaperatures
much less than 20°C.'§920 Dalhousie Springs alse
contains endemic snails {family| Hydoobiidae) and
amphinods.°
The cause of the unique morphology of the Dalhousic
Springs specimens (Le. genetic or envirtinenental) may be
solved by the rearing of young under conitollod conditions
The replication of this procedure with *pood" C
destructor would serve.as a centrol Without this further
evidenue, however, firm txonomic conclusions are nul
justified,
1 am indebted to Dr Sam Lake for his constructive
critigism and suggestions: My thanks also go 1 Dr bin
Bayly and Michael Barry for thet comments on this
manuscript.
' Sokal, A. (1986) Ph.D. Thesis, Department of Zoology,
Monash Lniversily, Victoria. (Linpublished),
q {in presay J. Invertebrate Taxenanyy,
3 Harris, C. R. (1985) In J. Greenslade, L. Joseph & A.
Reeves (eds) “South Australia’s Mound Springs”. (Nature
Conservation Society of South Australia Inc. Adelaide).
4 Riek, E. F. (1969) Aust. J. Zool. 17, 855-918.
5 Zar, + H. (1974) “Biostatistical Analysis” (Prentice-
Hall Inc. Englewood Cliffs, N.J.).
® Krieg, G. W. (1985) Dalhousie, South Australia,
1:250 000 Geological Series — Sheet SA/53-11 Explana-
tory Notes. (Geological Survey of South Australia),
personal communication,
8 5 Kolding, S. (1985) Marine Biology 89, 249-255.
9 Ponder, W. (1986) Zn P. De Deckker & W. D. Williams
(eds) “Limnology in Australia”. (CSIRO-Dr W. Junk
ip Publishers, Melbourne).
'" Hobbs, H. H. Jr. (1958) Q. J. Fla Acad. Sci. 21, 71-91.
ua (1969) Jn P. C, Holt (ed) “The Distributional!
History af the Biota of the Southern Appalachians. Pt 1
Invertebrates”, (Research Division Monograph 1. Virginia
Polytechnic Institute. Blacksburg, Virginia).
2 Williams, A. F. (1974) Sampling and measurement of
Great Artesian Basin, Mound Springs, South Australia,
209
Progress Report. 1. Dalhousie Springs. Department of
Mines, South Australia. Report 74/204.
'3 Lake, PS. & Sokol, A. (1986) The ecology of the
yabby Cherax destructor Clark (Crustacea: Decapoda:
Parastacidae) and its potential as a sentinel animal for
mercury and lead pollution, Australian Water Resources
Council, Technical Paper No. 87.
‘4 Faragher, R. A. (1983) Aust. J, Mar. Freshwater Res.
Is22! 407-417.
‘5 Wenner, A. M., Fusaro, C. & Oaten, A. (1972) Can. J.
Zool. 52, 1095-1106.
i 7 Clover, C. J. M. personal communication,
1, Zeidler, W. personal communication.
5 Wwantsoff, W. & Glover, C, J. M. (1974) Aust. Zool. 18,
88-98.
'9 Glover, C. J. M. (1982) In W. R. Barker & P. J. M.
Greenslade (eds) “Evolution of the flora and fauna of
arid Australia”, (Peacock Publications, Frewville, South
Australia.)
20 Merrick, J. R. & Schmida, G, E. (1984) Australian
Freshwater Fishes, (Griffin Press Limited, Adelaide).
ANTHONY SOKOL, Department of Zoology, Monash University, Clayton, Vic, 3168.
MIDDLE EOCENE AGE OF THE MEGAFOSSIL FLORA AT GOLDEN
GROVE, SOUTH AUSTRALIA: PRELIMINARY REPORT, AND
COMPARISON WITH THE MASLIN BAY FLORA
Summary
BRIEF COMMUNICATION
MIDDLE EOCENE AGE OF THE MEGAFOSSIL FLORA AT GOLDEN GROVE,
SOUTH AUSTRALIA: PRELIMINARY REPORT, AND COMPARISON WITH
THE MASLIN BAY FLORA
A clay lens exposed during mining operations in »
Monier Golden Grove sand pit contains abundant remains
of leaves, pollen and spores, Evaluation of all the Tossils
continues but preliminary results of the mezalosuils are
now available’.
This note presents the preliminary results of a
palynqological examination of two samples fram Golden
Grove and two others from the Jeal-bearing beds at Maslin
Ray2 4 The latter saniples were examined In order to
determine whether che two sites are correlative, A more
extensive treatment of the palynofloras from Golden Grove
will be given following analyses of borehole saraples of
the glay leds obtained dunte a recent drilling programme
in the Golden Grove area’. Detailed records of the
palynofloras are held m the Biostratigraphy Bronch,
Department of Mines and Fnergy, South Australia.
All samples produced diverse: well preserved
palynotloras in which pollen of Nothofuyidites spp.,
Haloragacidites horrisii (Couper) Harris 1971, the
pymnosperms, the Preteaceae and spores: from the
cryptogams are very common. No one group, however,
is @ Very dominant part of the assemblage.as is usually
expected, say, of Hualeragacidites Aarrisii and
Nothojagidites spp. in Tertiary palynofloras. The samples
are alsa rich in their diversity of triporate, Iricolpate and
tricolporate pollen produced by the angiosperms. One
sumple from Golden Grove contained an unusually high
frequency of tricalporate pollen, in particular that assigned
ta Rhoipites sphaerica (Cookson) Pocknalt & Crosbie
1982. In general, however, the composition of the
assemblages from cact site is similar.
In southern Australia two palynological zones have been
erecyed for the Tertiary in the Gippsland and Souch
Ausiralian aveas (Fig. J) In the Harcis zonation,
palynofloras from Golden Grove and Maslin Bay can he
assigned to rhe R pachypolus Zone, This derermination
\5 made on the hasis of the presence of Norhafasidites
asperus (Cookson) Stover & Evans 1973 and N. falcaius
(Cookson) Stover & Evans 1973, the first occurrence of
which marks the base of the Zone, as well as (he common
occurrence of the zonal species together with frequent
Norhofagidites spp. and Proteacidites spp., particularly
P. Kepiensis Harris 1972, 2 reticulatus Cookson 1980 and
P symphyonemaides Cookson 1950°. Trifites tuberculi-
formis Cookson 1947 and Matonispuritey ornamentatix
(Cookson) Stover & Partridge 1973 are rare, and THorites
magnificus Cookson 1950 is absent; The firsr occurrence
of the latter species is used to mark the top of the P
pachypolus Zone,
The palynotioras are also correlatives of the lower
Nothofozidites asperus Zone in the Gippsland sequences
(Fig. 1), thus supporting the conclusion drawn above. The
base of fhe Zone is marked by the first appearance of a
number of species which are present in the palynofloras
inchiding: Foventriletes bajteus Partridge 1973, Gerimerri-
volporites gesfus Partridge 1973, Gathanipallis hassensis
Staver 1973, Nothofasidites axperus (Cookson) Stover &
Evans 1973, N._faledtus (Cookson) Stover & Evans 1973,
PALYNOLOGICAL
ZONES
GIPPSLAND BASIN | SGUTH AUSTRALIA
(After 9,10) Aller Bt)
PALYNOLOGICAL
ZONES
Unnamed urt
Triporapaliensies
bellus
Cyolheocidiles
onnulate
Proreacid|tes
fubercuiatus
Nothotagidites
Prateacidjles
pachypolus
OSPErUs
Praleacidites
confragosus
Prot asperopolus
Malvocipe iis
diversus arthateichus
Gombierina
edwards |
Lygistepollenites
balmei
Tricaiplres
Jongus
Tricolpires
longus
Fig. |. Tertiary palynological zones in southern Anstralia,
N, vansteenisi (Cookson) Stover & Byans 1973, Peripore-
pollenites vesicus Partridge 1973, Rheipites unzurium
(Patttidge) Pocknall & Crosbie 1982, Augulatisporites
trophus Partridge 1973, Tricolpites simarus Partridge 1973,
T thomasii Cookson & Pike 1954 and Tricolporites leuros
Partridge 197390.
A number of species whose upper range terminates
within theJower NV, asperus Zone are also present including
22
Anacolosidites acutullus Cookson & Pike 1954, A.
Iuteaides Cookson-& Pike 1954, A. sectus Partridge 1973,
Dryptopollenires semilunatus Stover 1973, Proteacidijes
dlveolatus Stover 1973, PB asperopolus Stover & Evans
1973, P. incurvaius Cookson 1950, PB kopiensis Harris
1972, PF. pachypolus Cookson & Pike 1954, P tenuiexinus
Stover 1973, Palycolpites reticulatus Couper 196),
Rhoipites sphaerica and Verrucosisporites cristatus Part-
ridge 1973.
Palynofloras previously recovered from the Maslin Bay
leaf beds were believed to have been correlative with the
older Proteacicdites confragosus Zone, although in the light
of Jater work they were reassigned to the Profeucidites
pachypolus Zone®'*, The data presented in the present
note not only support the latter conclusion bur alsa show
that the palynofloras from Maslin Bay and Golden Grove
are correlative. Thus the Golden Grove and Maslin Bay
clay lenses and plant megafossils are Middle Eocene in
age, as are the fossil floras from the Maslin Bay site. The
palynological zonations and the age determination indicate
that the sand deposit at Golden Grove is a correlative of
the North Maslin Sands.
The palynofloras found at Golden Grove appeat to be
more diverse than the macrofloral assemblages’. At least
134 taxa can be recognized in the palynoflora, of which
115 can be ascribed to established form yenera or species,
and a further 19 species of unknown affinity. This is to
be expected, however, because the pollen and spores. are
IChristephel, D. C, & Greenwood, D, R, (1987) Trans.
R, Soc. 5. Aust. JIL, 155-162.
2Blackburn, D. T. (1981) Alcheringa 5, 9-28.
3Chrigtuphel, B.C. (1981) Jn Keast, A. (Ed.), “Ecological
biogeography of Australia” (W. Junk, The Hague).
& Biackburn, D. T. (1978) Alcheringa 2,
4,
31h-319.
SLange, R. T. (1970) N. Jb, Geol. Palaont, Mh. 8,
486-490.
'bMeGowran, B., Harris, W. K. & Lindsay, J. M. (1970)
N. Jb. Geol. Palaont. Mh, 8, 481-485,
7Pept. Min. Ener, §. Aust, (1986) Ja Miner, Ind. Quart.
43, 30-31.
representative of the regional vegetation and may have:
been transported considerable distances to the site. The
macrofossils, on the other hand, were derived in close
proximity to the site of deposition and provide evidence
of the local vegetation, Moreover, the sediments in which
the macrofossils occur are very finely laminated clay and
sill with no current bedding to indicate that the leaves
could have been brought from some distance away by
streams.
Pollen and spores from a number of plant geneva
recorded at Golden Grove are well represented in the
palynofloras, Proteaceous pollen is common and diverse,
including pollen of Banksia Linnaeus f, and Grevillea R.
Brown ex R.. A, Salishury, as.is pollen from the family
Podocarpacege, inchiding three species of Pkylloctadus
L. C. Rich, one of Dacrydiun Solander, three of
Podocarpus L’Herit ex’ Pers, and one Microcachrys F.
Hooker. The Myrtaceae are common and at least four
species of pollen can be recognized, including pollen with
affinities ta Euca/yvprus L’Herit. Although spores of the
eryptogams form only small proportion of the assemblage,
they are diverse and include two specics of Lygodiym
Linnaeus f. type spores and two others with probable
affinities to the genus.
Lthank J. M. Lindsay, W. V. Preiss, W. K. Harris and
C, B, Foster for their comments on the manusertipr,
This disenssion is published with the permission of the
Nitector-General, Department of Mines & Energy.
‘Harris, W. K. (1971) Jr Wopfner, H, & Douglas, J, G.
(Eds). The Orway Basin of Southeastern Australia, Spec,
Bull. geol. ‘Survs. 5. Aust. & Vict. 67-67.
*Stover, L. E. & Partridge, A. D. (1973) Proc. R. Soc.
Vict. 85, 237-286.
0 & -——— (1982) Palynology 6, 69-95.
il Truswell, £. M..& Harris, W. K. (1982) Jn Barker, W.
S. & Greenslade, P. f, M, (Eds), “Evolution of the flora
and fauna of arid Australia” (Peacock Publications,
Adelaide) pp. 67-76.
'2Harris, W, &. (1985), Ja Lindsay, J. M, (Fd,)
“Stratigraphy, Palacontology, Malacolopy. Papers in
honour of Dr Nell Ludbrook”. Spec. Publ. Dept. Mir.
Enet, 5, Aust, 5, 133-144.
NEVILLE F, ALLEY, Department of Mines and Pnergy, South Australia, P-O. Box 151, Eastwood, 5. Aust. 5003.
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